Mémoires du Muséum national d'histoire naturelle Série A Zoologie

cm :m

ZOOLOGIE

TOME 150

1991

Coordonné par

Alain CROSNIER
& Philippe Bouchet

MÉMOIRES
DU MUSÉUM
NATIONAL
D’HISTOIRE
NATURELLE

Résultats des Campagnes MUSORSTOM

Volume 7

MÉMOIRES DU MUSÉUM NATIONAL D’HISTOIRE NATURELLE

Directeur de la publication : Philippe Bouchet
Rédacteurs (Editors) : P. Bouchet, A. Dubois, C. Erard
Secrétariat : Bernadette Charles
Conception graphique : Alain Defilippi
Rédaction : 57, rue Cuvier
75005 Paris

Les Mémoires du Muséum national d’Histoire natu¬
relle publient des travaux originaux majeurs (100
pages et plus) dans les domaines suivants : Zoologie
(série A), Botanique (série B). Sciences de la Terre
(série C). Les auteurs sont invités, pour toutes les
questions éditoriales, à prendre contact avec le direc¬
teur de la publication.

Vente en France
(uniquement)
Éditions du Muséum
Lionel Gauthier
38, rue Geoffroy St-Hilaire
75005 Paris
Tél. : (1) 40-79-37-00
Telex musnahn 202641 F
Fax : 40-79-34-84

Parution et prix irréguliers. Les ordres permanents
d'achat et les commandes de volumes séparés sont
reçus par le service de vente. Catalogue sur demande.
Une liste des derniers titres parus figure en page 3 de
couverture.

Mémoires du Muséum national d’Histoire naturelle
publishes major original contributions (100 pages and
over) in three different sériés : Zoology (série A),
Botany (série B), Earth Sciences (série C). Prospective
authors should contact the Editor. Manuscripts in
French and English will be considered.

Sales Office
(France excluded)
Universal Book Services
Dr. W. Backhuys
Warmonderweg 80
2341 KZ Oegstgeest
The Netherlands
Tel. : (71) 17 02 08
Fax : (71) 17 18 56

Volumes are published at irregular intervals, and at
irregular prices. Standing orders and orders for single
volumes should be directed to the Sales Offices. Free
price list and catalogue on request. Recently published
memoirs are listed on page 3 of the cover.

The Muséum national d'Histoire naturelle also
publishes a Bulletin.

Source : MNHN, Paris

Source : MNHN, Paris

Résultats des Campagnes Musorstom
Volumes déjà parus :

Volume 1
Volume 2
Volume 3
Volume 4
Volume 5
Volume 6

Volume 7

Mém. Orstom, 91 : 1-558, 225 fig., 39 pl. (1981). ISBN : 2-7099-0578-7.

Mém. Mus. natn. Hist. nat. Paris, (A), 133 : 1-525, 126 fig., 37 pl. (1986). ISBN : 2-85653-136-9.

Mém. Mus. natn. Hist. nat. Paris, (A), 137 : 1-254, 82 fig., 9 pl. (1987). ISBN : 2-85653-141-5.

Mém. Mus. natn. Hist. nat. Paris, (A), 143 : 1-260. 103 fig., 23 pl. (1989). ISBN : 2-85653-150-4.

Mém. Mus. natn. Hist. nat. Paris. (A), 144 : 1-385, 128 fig., 35 pl. (1989). ISBN : 2-85653-164-4.

Mém. Mus. natn. Hist. nat. Paris, (A), 145 : 1-388, 190 fig., 4 pl. couleur (1990).
ISBN : 2-85653-171-7.

Mém. Mus. natn. Hist. nat. Paris, (A), 150 : 1-264, 587 fig., (1991). ISBN : 2-85653-180-6.

Source : MNHN, Paris

ésuUats des campagnes

SB

Volume 7

ISBN : 2-85653-180-6
ISSN : 0078-9747

© Éditions du Muséum national d’Histoire naturelle, Paris, 1991.

Source : MNHN, Paris

MÉMOIRES DU MUSÉUM NATIONAL D’HISTOIRE NATURELLE

SÉRIE A

ZOOLOGIE

TOME 150

Résultats des Campagnes MUSORSTOM

Volume 7

Coordonné par

Alain CROSNIER

Muséum national d’Histoire naturelle
Laboratoire de Zoologie, Arthropodes
61, rue Buffon
75005 Paris

&

Philippe Bouchet

Muséum national d’Histoire naturelle
Laboratoire de Biologie des Invertébrés marins et Malacologie
55, rue Buffon
75005 Paris

Publié avec le concours de I’orstom

ÉDITIONS
DU MUSÉUM
PARIS

1991

Source : MNHN, Paris

Source : MNHN, Paris

Sommaire

Contents

1. Mollusca Polyplacophora : Deep-water Chitons from New Caiedonia . 9

Pieter Kaas

2. Mollusca Bivalvia : Archibenthal Nuculidae off New Caiedonia . 29

Wim Bergmans

3. Mollusca Gastropoda : Seguenziidae from New Caiedonia and the Loyalty Islands 41

Bruce A. Marshall

4. Mollusca Gastropoda : Systematic position and révision of Haloceras Dali, 1889

(Caenogastropoda, Haloceratidae fam. nov.) . 111

Anders Waren & Philippe Bouchet

5. Mollusca Gastropoda : Four new rissoinine species (Rissoidae, Rissoininae) from deep water

in the New Caledonian région. 163

Willy J. Sleurs

6. Mollusca Gastropoda : Cypraeopsis superstes sp. nov., Pediculariinae relique du bathyal de

Nouvelle-Calédonie et de la Réunion. 179

Luc Dolin

7. Mollusca Gastropoda : On a collection of Nassariidae from New Caledonian waters . 187

Walter O. Cernohorsky

8. Mollusca Gastropoda : Eumitra récentes de la région néo-calédonienne et Charitodoron

fossiles de l’Oligocène supérieur d'Aquitaine (Mitridae). 205

Pierre Lozouet

9. Mollusca Gastropoda : The Typhinae (Muricidae) from the New Caledonian région with

description of five new species. 223

Roland Houart

10. Mollusca Gastropoda : Columbariform Gastropods of New Caiedonia. 243

M. G. Harasewych

Source : MNHN, Paris

Source : MNHN, Paris

5ULTATS DES CAMPAGNES MUSORSTOM, VOLUME 7 - RÉSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSUI

1

Mollusca Polyplacophora : Deep-water
Chitons from New Caledonia

Pieter KAAS

Nationaal Natuurhistorisch Muséum
P.O. Box 9517
2300 RA Leiden
The Netherlands

ABSTRACT

Five Frcnch deep-sea cruises made around New Caledonia
during lhe years 1985-1987 brought altogether 92 spccimens
of chitons, representing 10 species in 5 families ; 8 specics are

new to science. The new genus Vermichiton is described for a
small vermiform spccies ; this genus is compared with
Connexochiton Kaas, 1979.

RÉSUMÉ

Mollusca Polyplacophora : Chitons bathyaux et abyssaux de
Nouvelle-Calédonie.

Cinq campagnes françaises réalisées autour de la Nouvelle-
Calédonie, de 1985 à 1987, ont permis la récolte de 92

spécimens de chitons, appartenant à cinq familles et compre¬
nant dix espèces dont huit nouvelles. Le nouveau genre
Vermichiton est décrit pour une petite espèce vermiforme : ce
genre est comparé avec Connexochiton Kaas. 1979.

Kaas, P.. 1991. Mollusca Polyplacophora : Deep-water Chitons from New Caledonia. In : A. Crosnier & P. Bouchet (eds),
Résultats des Campagnes Musorstom, Volume 7. Mém. Mus. nain. Hist. nat.. (A), 150 : 9-27. Paris ISBN : 2-85653-180-6.

Publié le 20 mars 1991.

Source : MNHN, Paris

10

PIETER KAAS

INTRODUCTION

During the years 1985-1987 several deep-sea
cruises were made around New Caledonia. In
1985 the Biocai. cruise, on the research vessel
“ Jean Charcot ", procured 43 specimens of
Polyplacophora from 8 stations S and SE of
New Caledonia, in depths varying from 235 to
2 340 m. They belong to 6 species, 4 of them new
to science. Later in 1985 the Musorstom 4
cruise, on the research vessel “ Vauban ", hauled
36 specimens of chitons from 12 stations NW
and SE of New Caledonia in depth from 75 to
720 m, representing 6 species, 4 new to science,
one had also been procured by the Biocal cruise.
In 1986 the Chalcal 2 cruise added 11 more
specimens, belonging to 3 species, two already
found by the aforementioned cruises and one
new. The same year the Smib 2 and the Smib 3
cruises, on the “ Vauban ", added two more
specimens of species also found by the foregoing
cruises. Altogether 92 specimens were captured,
representing 10 species, 8 of which are new to
science.

B. Richer de Forges (1990) wrote a narrative
of the cruises and provided a list of stations.

Thanks are due to Dr Philippe Bogchet of the
Muséum national d’Histoire naturelle, Paris,
who participated in the Biocal, Musorstom 4
and Chalcal 2 cruises, for afîording the author
an opportunity to study and describe the mate-
rial.

List of abbreviations :
ams = Australian Muséum, Sydney.
mnnh = Muséum national d'Histoire naturelle,
Paris.

nmnz = National Muséum of New Zealand,
Wellington.

rmnh = Rijksmuseum van Natuurlijke Historié,
torie, Leiden. Now Nationaal
Natuurhistorisch Muséum.
vb = Private collection of R. A. Van Belle,
Sint-Niklaas, Belgium.

SYSTEMATIC ACCOUNT

Order NEOLORICATA
Suborder LEPIDOPLEURINA
Family LEPTOCHITONIDAE
Genus LEPTOCHITON Gray, 1847
Subgenus LEPTOCH1TON s.s.

Leptochiton (L.) belknapi Dali, 1878

Leptochiton belknapi Dali. 1878 : 1. Kaas &
Van Belle, 1987 : 23, fig. 10 (bibliography and
synonymy). Kaas, 1990 : 176.

Material examined. New Caledonia. Biocal :
stn CP 05. 2116' S. 166"44' E, 2 340 m, 11 .VIII. 1985 :
I spm. Stn DW 33, 23“10'S, 167"10'E, 675-
680 m. 29.VIII. 1985 : I spm. Stn CP 72, 22°10' S,
I67‘’33'E. 2 100-2 110 m, 04.IX.I985 : 2 spms.

Leptochiton (L.) perscitus sp. nov.

Figs 1-12

Material examined. New Caledonia. Musor¬
stom 4 : stn DW 149. 19 Ü 08'S, I63"23' E. 155 m.
14.IX.1985 : 18 spms, paratypes (mniin/12, rmnh
9264/2, vu 2983/2. nmnz/1). Stn DW 150, 19‘'07' S,
163“22' E. 110 m, 14.IX.1985 : 3 spms. holotypc and
disarticulated paratypes (mnhn).

Diagnosis. — Animal c. 2 mm long, modera-
tely elevated, subcarinated, valves nol beaked.

Source : MNHN, Paris

MOLLUSCA POLYPLACOPHORA : DEEP-WATER CHITONS

Map showing stations of the cruiscs Biocal, 1985 (numbers < 100) and Musorstom 4, 1985 (numbers > 100).

Source : MNHN, Paris

12

PIETER KAAS

Figs 1-12. Leptochiton (L.) perscitus sp. nov. : 1. valve I. dorsal view. * 46. 2. valve II. dorsal view, x 46. 3, valve

VII, dorsal view, x 46. 4. ici. rosirai view, x 23. 5. valve VIII. dorsal view, x 46. 6. /</.. laierul view, x 46

7. different types of dorsal girdle scalcs. x 460. 8. marginal spiculé, x 460. 9, suturai spiculés. < 460. 10

ventral sca le. x 460. 11. central and firsl latéral radula teeth, • 460. 12. different views oflicad of major latéral

looth. x 460. 1-12. paratype from Mijsorstom 4 eruisc. stn DW 150.

latéral areas little raised, tail valve short, mucro
post-median. End valves and latéral areas weakly
quincuncially granulated, central and antemu-
cronal areas with larger, roundish granules in
longitudinal sériés. Girdle with linely striated
scales, at the sutures oceasionally small bunches
of spiculés. Dental cap of major latéral radula
tooth bicuspid.

Description. Animal very small. the largest
2.1 x 1.8 mm (curled). at most 2.5 mm long
when stretched. oval, subcarinated, side slopes

slightly convex, moderately elevated (dorsal élé¬
vation c. 0 . 40 ), valves not beaked.

Valve I somewhat less than semicircular, quin¬
cuncially sculptured with weakly pronounced.
rounded granules. Intermediate valves rectangu-
lar. anterior margin convex in valve II, straight
to a little concave in the others, posterior margin
almost straight, often damaged as the valves are
very thin and brittle. Latéral areas weakly raised,
quincuncially granulated like head valve, central
areas with longitudinal rows of well pronounced.
round granules, more weakly developed on the

Source : MNHN, Paris

MOLLUSCA POLYPLACOPIIORA : DEEP-WATER CHITONS

13

jugum. Apophyses small, short, triangular to
trapezoid, widely spaced, jugal sinus almost fiat.
Tail valve short, length less than half ils width,
the mucro post-median, not swollen, posterior
slope steep, concave. Antemucronal area sculp-
tured like the central areas, postmucronal area
like the head valve and latéral areas.

Girdle white, narrow, densely paved with
finely longitudinally striated seules of various
forms and sizes, but always at least twice as long
as wide, with more or less rounded top, up to
75 tzm long, 30 [xtn wide. Occasionally, small
bunches of straight, slender, smooth spiculés
occur at the sutures, varying in length from 66-
100 |xm. Marginal spiculés sharply pointed, lon¬
gitudinally grooved, 60 x 10 jzm. Ventral side
of girdle paved with radiating rows of fiat scales,
c. 55 x 25 |zm, distally narrowing to a blunt,
grooved point.

Central tooth of radula narrow, slightly pinch-
ed in the distal half, with a rounded blade, first
latéral weakly curved inwardly, with a small
blade, dental cap of major laterals with a sharply
bent, pointed main cusp and a short, small
external cusp.

Etymology. The Latin perscitus = very
fine, in relation to the delicacy of the shell.

Discussion. Until now species of Leptochi-
ton hâve not been reported from New Caledonia.
L. perscitus resembles L. norfolcensis (Hedley
& Hull, 1912) from Lord Howe and Norfolk
Islands and its supposed subspecies subtropicales
(Iredale, 1914) from the Kermadec Islands,
which, however, grows much larger, to 6 mm
long and has much shorter, sparsely but strongly
ribbed dorsal girdle scales.

Leptochiton (L.) vuubam sp. nov.

Figs 13-23

Matiîrial examine». New Caledonia. Musor-
stom 4 : stn DC 168, 18"48' S, 163°H'E, 720 m,
16.IX. 1985 : 1 spm, holotype (MNHN).

Diagnosis. Animal c. 2 x 1 cm, elongate
oval, moderately elevated, carinated. Colour
white. Valves not beaked, latéral areas not
raised, mucro almost central. Apophyses ischno-
chitonoid, rather wide. End valves and latéral
areas with small, elevated pustules arranged in
curved sériés, continuing on central areas, where

they are longitudinally arranged and less ele¬
vated. Girdle with elongate, striated spiculés.
Major latéral teeth of radula with bicuspid
dental cap.

Description. — The holotype measures 19.8
x 9.9 mm (now disarticulated. slides of perino-
tum and radula). Dorsal élévation 0.44. which
is moderate, the back carinated, side slopes
straight. Colour of tegmentum and girdle white.

Head valve less than semicircular, anterior
slope straight, posterior margin widely V-shaped.
Intermediate valves broadly rectangular. side
margins only little convex, anterior margin
slightly convex, except for the jugal sinus which
is decidedly concave ; posterior margin straight,
the apex not or hardly projecting. Latéral areas
not raised, marked only by the différence in
orientation of the sculpture, consisting of chains
of well raised, small, roundish pustules, arranged
in curved sériés in two directions, forming a
neatly quincuncial pattern. The sériés continue
across the central area in parallel longitudinal
Unes, beeoming less pronounced, almost obsolète
on the jugum. Tail valve slightly narrower than
head valve, the length about 3/5 of the breadth,
the mucro about central, not swollen, posterior
slope concave directly behind the mucro. Ante¬
mucronal area sculptured like the central areas,
postmucronal area and head valve like the latéral
areas.

Articulamentum thin, white. apophyses rather
long, widely triangular in the intermediate
valves, more or less trapezoid in the tail valve,
ischnochitonoid, jugal sinus about one quarter of
the valves’s width.

Girdle narrow, covered with elongate, sharply
pointed, longitudinally grooved spiculés, 88 x
24 jj.m on mid-girdle, up to 160 x 30 |j.m at the
inner margin. The cuticula of the girdle bridges
bears long, glassy, slender and smooth needles,
up to 300 x 20 |zm. Ventral scales ovoid with
pointed top, slightly striated, c. 28 x 24 izm.

Radula with a short and relatively wide central
tooth, slightly bulbous proximally, with a nar¬
row, straight blade ; first laterals widening dis¬
tally, ending in a shallow sinus, without a blade :
major laterals with a strong, curved, bicuspid
dental cap, the cusps pointed, the central one
much larger than the outer one.

Gills merobranchial abanal : c. 18 ctenidia per
side.

Source : MNHN, Paris

14

PIETER KAAS

Figs 13-23. Leptochiton (L.) vaubani sp. nov. : 13. valve I, dorsal view, x 9.6. — 14. valve IV, dorsal view, x 9.6. 15.

id.. rostral view, x 4.8. 16. valve VIII, dorsal view, x 9.6.- 17, id., latéral view, x 9.6. 18. dorsal girdlc spiculés

along valve margin, x 200. 19, id., from mid-girdle, x 200. 20, needles from girdle bridges, x 200. 21. ventral

spiculés, x 200. 22, central and first latéral radula tecth, x |00. 23, head of major latéral tooth

x 100. — 13-23, holotype.

Etymology. - After the research vessel
“ Vauban

Discussion. — L . vaubani does not resemble
any known species of the genus. It is unique in
the possession of strongly developed, ischnochi-
tonoid apophyses, in the spiculose girdle and the
needle-bearing girdle-bridges.

Leptochiton (L.) sp. indet.

Material examiner. New Caledonia. Biocal :
stn CP 05, 2ri6'S, 166°44' E, 2 340 m, 11,VIII. 1985 :
2 spms, too small to identify. — Stn DW 51, 23°05' S,
167°45' E, 680-700 m, 03.IX.I986 : 4 spms. smashed,
unidentifiable.

Source : MNHN, Paris

MOLLUSCA POLYPLACOPHORA : DEEP-WATER CHITONS

15

Suborder 1SCHNOCH1TONINA
Family 1SCHNOCH1TONIDAE
Subfamily ISCHNOCHITONINAE

Genus VERMICHITON gen. nov.

Diagnosis. Animal small, narrowly elon-
gate, more than four times longer than wide,
highly elevated, carinated, apophyses connected
by a jugal plate, articulamentum with many/1/

many slits, girdle closely beset with bluntly
pointed, juxtaposed spiculés, directed towards
the outer margin.

Type : V. vermiculus sp. nov.

Vermichiton vermiculus sp. nov.
Figs 24-36

Material examined. New Caledonia. Biocal :
stn DW 46, 22"53'S, 167°17'E, 570-610 m, 30.VIII.
1985 : 1 spm, partly disarticulated. holotype (mnhn).

Diagnosis. Animal small, c. 5 mm long,
1 mm wide, valves highly elevated, carinated,
hardly or not beaked. End valves and latéral
areas pustulose, central areas finely longitudi-
nally punctate-striate, apophyses connected by a
jugal plate, slit formula of insertion plates
many/1/many. Girdle with callochitonoid spi¬
culés.

Description. Animal small, narrowly elon-
gate, 5.12 x 1.36 mm, highly elevated (dorsal
élévation 0.63), sharply carinated, side slopes
almost straight, valves not beaked, apices hardly
indicated, latéral areas raised. Girdle relati-
vely wide, covered with callochitonoid spiculés.
Colour of valves and girdle white.

Head valve little more than semicircular, ante-
rior slope steep, straight, posterior margin bay-
like ; intermediate valves relatively long, not or
slightly beaked, latéral areas clearly indicated,
raised, anterior margin almost straight, convex
in valve II. which is longer than the others,
side margins straight, side slopes steep, almost
straight. Posterior valve as long as wide, the
mucro central, slightly directed forwardly, poste¬

rior slope convex. Sculpture consisting of relati¬
vely large, raised, round, quincuncially arranged
pustules on head valve, latéral areas of interme¬
diate valves and postmucronal area of tail valve ;
central and antemucronal areas finely, longitudi¬
nal^ punctate-striate. obsolète on the jugum.
End valves and latéral areas with a few concen-
tric growth marks.

Articulamentum well developed. white, pro-
duced forwardly in valves II-V1II as to form a
relatively long jugal plate, connecting the apo¬
physes. As a resuit the jugal sinus is very
shallow. Valve I with 14 slits, the insertion
teeth smooth, eaves solid ; intermediate valves
with 1-1 short slits, valve VIII with 16 slits, the
teeth very short and blunt. There are no slit-rays.

Girdle relatively wide, directed downward.
appearing narrow when viewed from above,
dorsally covered with juxtaposed, bluntly point¬
ed, elongate-ovoid, white spiculés, neatly ar¬
ranged in quincunx, c. 150 x 40 um. There is a
marginal fringe of two kinds of small, pointed
spiculés : white, smooth, slender ones, c. 120 x
15 jxm, and stout, longitudinaliy grooved ones,
120 x 30 jxm. Ventral spiculés arranged in ra-
diating rows, the basal half distinctly striated.
distally abruptly narrowing to a needle-like
point, 90 x 16 ^m.

Animal white, with a rounded head about the

Source : MNHN, Paris

16

PIETER KAAS

FifiS 24-36. VermichUon gen. nov. vermiculus sp. nov. : 24-26, dorsal, ventral and latéral view respectively, x 18.75. 27-

28. valve I. dorsal and anterior view respectively. x 18.75. 29-30. valve II, dorsal and ’rostral view respectively,

x 18.75. 31-33, valve III. dorsal, ventral and latéral view respectively, x 18.75. 34. two types or marginai

spiculés, x 150. 35. dorsal girdle spiculés in situ, x 75. 36. ventral scalcs. x 300. 24-36, holotype.

length of the foot, which is narrow, the width at
most 1/7 of the length. Gills merobranchial,
abanal, about 20 ctenidia per side.

Radula not examined.

Etymology. — From the Latin vermis =
worm. so worm-like chiton ; vermiculus= dimi-
nutive of vermis, meaning little worm, relating to
the scantiness and worm-like appearance of the
animal.

Discussion. — This remarkable species shows
some affinilies to the genus Callochilon Gray,

1847, especially in the slitting of the insertion
plates and the presence of a jugal plate in the
valves II-V1II. The texture of the tegmentum,
however, along with the elongate shape of the
animal and the absence of extrapigmentary eyes
in the shell plates, are of a more ischnochitonoid
character. That is why the new genus Vermichi-
ton has been created for this unique species,
which should be classified with Connexochiton
Kaas, 1979.

Source : MNHN, Paris

MOLLUSCA POLYPLACOPHORA : DEEP-WATER CHITONS

17

Genus COISNEXOCH1TON Kaas, 1979

Connexochiton discernihilis sp. nov.

Figs 37-49

Material examined. — New Caledonia. Biocal :
stn DW 44, 22"47'S, 167” 14' E. 440-450 m, 30.V11I.
1985 : 17 spms, MNHN/holotype + 10 paratypes;
rmnh 9265/2 paratypes ; vb 2984/2 paratypes ; ams/
1 paratype; nmnz/1 paratype.

Diagnosis. — Animal small, elongate oval, up
to 7.5 x 3.5 mm, white, highly elevated, side
slopes straight, back carinated.

Latéral areas raised, with 5-7 radial sulci
crossed by numerous fine, concentric Unes. Cen¬
tral areas with a vague pattern of dépréssions,
parallel to the diagonal ridges, crossed by con¬
centric striae. Head valve and postmucronal area
sculptured like latéral areas. Apophyses connect-
ed by a jugal plate, showing a small sinus in the
médian line of valves III-VII. Girdle narrow,
covered with small, slriated scales.

Description. Animal elongate oval, twice
as long as wide, small, the largest c. 7.5 x
3.5 mm, highly elevated (dorsal élévation c. 0.58),
side slopes straight, the back sharply carinated.
Valves slightly though sharply beaked. Colour of
tegmentum and girdle white.

Head valve semicircular, posterior margin
widely V-shaped, anterior slope steep, straight.
Valve II longer than the others, forwardly pro-
duced in the jugal région, the anterior margin
concave at both sides of the jugum. Valves III-
VII transversely rectangular, anterior and poste¬
rior margins almost straight, but for the apex,
forming a small but well marked beak. Side
margins a little convex. Latéral areas slightly
raised. Posterior valve crescent-shaped, more
than twice as wide as long, anterior margin
barely concave, mucro not prominent, central,
posterior slope rather fiat, only little excavated
directly behind the mucro. Sculpture weakly
pronounced, consisting of 6-8 fine radial grooves
in the latéral areas, 25-30 in the head valve, less
in the postmucronal area of the tail valve,
separating weakly granulose riblets and crossed
by numerous fine, concentric fines. Central areas

vaguely sculptured with a sériés of wavy déprés¬
sions parallel to the diagonal ridges, crossed by
concentric striae, the jugum practically smooth.

Articulamentum well developed, forming a
rather long jugal plate between the apophyses,
dorsally with a few striations, in valves III-VII
with a very small, narrow médian sinus. Slit
formula of insertion plates 9/1/8, teeth sharp,
slit rays distinct, eaves solid.

Girdle rather narrow, dorsally covered with
small, rather elongate, curved. imbricating scales,
the base elliptical. 72 [un wide, 80 (un high
on mid-girdle, up to 130 um along the valves’
edges. They are sculptured with about 18 paral¬
lel. longitudinal riblets. as wide as the finely
latticed interstices ; shortly before reaching the
rounded top of the scale the riblets break into a
reticulate pattern. Ventral scales elongate rectan¬
gular, distally rounded. 56 x 10 u.m on mid-
girdle. Marginal spiculés spindle-shaped, point-
ed, 80 x 16 |/m, with a few longitudinal riblets.

Central tooth of radula relatively wide, some-
what pinched in the middle, acutely widening in
the basal part, with a narrow, straight blade ;
niinor laterals narrow, elongate, the distal edge
wider, bilobed, wilhout a blade ; major laterals
with a single, strongly curved, sharply pointed
cusp.

Gills merobranchial, adanal with interspace ;
c. 13 ctenidia per side.

Etymology. The Latin adjective discernihilis
means discernible, as the species is easy to
distinguish.

Discussion. - This is the first Connexochiton
described from the Pacific Océan, the three
previously reported species are found in the deep
water of the Atlantic, viz. C. platynomenus Kaas,
1979, the type of the genus, from the eastern side
(Brittany to Morocco and in the Mediterranean
Sea), C. mordrai (Righi. 1973) and C. brom-
leyi (Ferreira, 1985) from the tropical western
side. C. discernihilis is easily recognizable by the
absence of a pustulose tegmental sculpture and
by its highly arched, sharply carinated shape.

Source : MNHN, Paris

18

PIETER K AA S

Figs 37-49. Connexochiton dbcernibilis sp. nov. : 37, complote spccimcn, dorsal view, x 6.6. 38. valve I. dorsal view

x 20.5. 39, valve II, dorsal view, x 20.5. 40, valve III, dorsal view, x 20.5. 41, id„ rosirai view, x 10. 42

valve VIII. dorsal view, x 20.5. 43, «/., latéral view, x 20.5. 44. dorsal scalc from mid-girdle, dorsal view

x 415. - 45. id., from inner margin. ventral view, x 205. 46. marginal spiculé, x 415. 47. ventral scalc

x 415. 48, central and first latéral radula teeth, x 415. 49, head of major latéral tooth, x 415. 37-49

paratypes (mnhn).

Source : MNHN, Paris

MOLLUSCA POLYPLACOPHORA : DF.EP-WATER CHITONS

19

Genus ISCHMOCHITOIW Gray, 1847
Subgenus STENOSEMVS von MiddendorfT, 1847

Ischnochiton (Stenosemus) délicat us sp. nov.

Figs 50-56

Material examined. — New Caledonia. Musor-
stom 4 : stn CP 167, 18°36'S, 163°06' E, 575 m,
16.IX.1985 : 1 spm, holotype (mnhn). — Stn DW 221,
22°59'S. 167-37'E, 515-560 m. 29.IX.1985 : 1 spm.
incomplète, paratype (mnhn).

Diagnosis. — Animal of small to moderate
size, elongate oval, more lhan twice as long as
wide, rather elevated, valves slightly beaked,
latéral areas little raised. Tegmentum white,
finely granulated ; head valve, latéral and post-
mucronal areas with many weak radial grooves,
crossed by fine, close-set concentric fines, central
and antemucronal areas with many parallel
longitudinal chains of very fine granules, obso¬
lète on the jugum ; mucro of tail valve slightly
antemedian. Girdle with short, globular, almost
smooth seules. Dental cap of major latéral radula
tooth with a single cusp.

Description. Animal white, rather elongate
oval, 17.5 x 7.0 mm. highly arched (dorsal élé¬
vation 0.47), carinated, side slopes weakly con-
vex. Intermediate valves of about equal width,
only slightly beaked, latéral areas little raised.
Tegmentum granular, girdle narrow, scaly.

Head valve semicircular, posterior margin
widely V-shaped, with a small notch at the apex,
anterior slope straight, tegmentum granular, the
granules transversely oval, arranged in radiating
rows, separated by narrow, shallow grooves in
the lower half. Intermediate valves rectangular,
front marging nearly straight, except in valve II,
which is forwardly produced, strongly convex
between the apophyses, hind margin straight,
with a small, hardly protruding apex. Side
margins little convex. Latéral areas well marked,
weakly raised, sculptured like head valve, with
c. 10 shallow grooves ; central area with 30 or
more longitudinal rows of roundish, separated,
little elevated granules on the pleural sides,
obsolète on the jugum. Tail valve with the mucro

a little anterior, not swollen. the posterior slope
straight, with only a slight excavation directly
behind the mucro. Antemucronal area sculptur¬
ed like central areas, postmucronal area like
head valve.

Articulamenlum well developed, white, apo¬
physes moderately wide, broadly rectangular
with rounded top, connected by a very short
jugal plate, hardly projecting beyond the tegmen¬
tum. Insertion plates rather short, smooth-edged
in valves I-VIl, finely toothed in VIII. Slit-for-
mula 7/1/7, slit rays distinct, eaves narrow, solid.

50-56. holotype.

Source : MNHN, Paris

20

PIETER KAAS

Girdle rather narrow, partly folded under,
regularly paved with dorsally globular, almost
smooth to obsoletely striated, ventrally concave
scales, up to 110 x 120 pm on mid-girdle,
rapidly decreasing in size towards the outer
margin, which bears a short fringe of conical,
weakly ribbed spiculés, up to 100 ixm long, 24 jxm
thick at the base. Ventral side of girdle covered
with radial rows of rectangular scales. slightly
emarginate proximally. rounded distally, 90 x
25 ,um on mid-girdle, shorter towards the outer
margin.

Radula 7.3 mm long, which is c. 40 percent of
the length of the body, with 28 rows of mature
teeth. Central tooth bulged in the proximal half,
with a weakly bilobed blade, first laterals of
about equal length, distally with small projec¬
tions in- and out-wardly, dental cap of major
latéral tooth with a single, sharply pointed,
strongly bent cusp.

Gills merobranchial, adanal with interspace ;
c. 22 clenidia per side.

Etymology. The Latin adjective delicatus
means fine of texture.

Discussion. — Of the eight hitherlo described
species of the subgenus Stenosemus only L. (S.)
substriatus Kaas & Van Belle, 1989 from the
Cape Verde Archipelago, bears a slight resem-
blance to the new species, although it differs
significantly in ils much smaller size, in the
absence of radial grooves in end valves and
latéral areas, in the decidedly ribbed girdle
scales and in the bicuspid dental cap of the major
latéral radula tooth.

Ischnochiton (Stenosemus) robustus sp. nov.

Figs 57-67

Material examined. New Calcdonia. Biocal :
stn DW 44. 22°47' S. I67"14' E. 440-450 m, 30.V1II.
1985 : 5 spms, paratypes, totally disarticulated (mnhn/
2, AMS/ 1, NMNZ/1. vb 2985/1).

Musorstom 4 : stn CP 193, 18°56'S, I63°23'E,
415 m. 19.IX.1985 : 2 spms. smashed. paratypes
(mnhn). Stn CP 194. 18°53'S. 163°22'E, 545 m.
19.IX. 1985 : 1 spm. paratype (mnhn). Stn DW 221,
22 U 59'S. 167"37' E. 535-560 m, 29.IX.1985 : 2 spms.
holotype (mnhn) and paratype (rmnh 9266). — Stn
DW 230. 22°57' S. I67°12' E. 390-420 m, 30.IX.1985 :

1 spm, paratype (mnhn).

Smib 2 : stn DW 10 : 22°55' S. 167°16' E. 490-495 m.
I8.IX.1986 : 1 spm dry, paratype (mnhn).

Diagnosis. Animal of moderate size, elon-
gate oval, twice as long as wide, c. 30 x 15 mm,
rather highly elevated, carinated, valves not
beaked, latéral areas weakly raised, obsoletely
radially ribbed, central areas with close-set,
forwardly converging sulci. Head valve and
postmucronal areas of tail valve sculptured like
latéral areas. Colour white. Girdle with curved,
bulbose, finely striated scales. Head of major
latéral radula tooth unicuspid.

Description. Animal elongate oval, holo¬
type 29.8 x 14.8 mm, highly elevated (dorsal
élévation 0.50), carinated. side slopes straight.
Valves of equal width, the intermediate ones
truncated at the outer margins, not beaked.

Head valve semicircular, posterior margin
widely V-shaped with a rounded apical notch,
anterior slope steep, straight, sculptured with
c. 40 obsolète radial grooves. Intermediate valves
rectangular. about twice as wide as long, front
margin somewhat sinuate, with the jugal sinus
slightly convex. the pleural parts concave, form-
ing a sharp angle with the side margins ; poste¬
rior margin straight, the apex not projecting,
hardly discernible. Latéral areas well marked,
only slightly raised, with 6-8 obsolète radial
grooves, crossed by numerous fine growth fines,
especially towards the outer margin. Central area
with c. 30 forwardly converging, well marked
sulci on either side, hardly narrower than the fiat
riblets in between. Tail valve short, outer mar¬
gin 1/3 of a circle, front margin straight, but for
a slight projection in the jugal sinus, mucro
almost central, slightly swollen, postmucronal
slope concave. Antemucronal area sculptured
like the central areas, postmucronal area like
the head valve and latéral areas.

Articulamentum well developed, white, apo¬
physes rather long and wide, bluntly triangular
in intermediate valves, trapezoid in tail valve,
always connected by a narrow but distinct,
dorsally striated jugal plate ; sinus straight to
somewhat convex. Insertion plates with 10/1/10-
12 inequidistant slits, slit-rays distinct ; eaves
solid.

Girdle moderately wide, paved with curbed,
dorsally bulbose, finely striated scales, c. 100 pm
wide, up to 140 (xm high on mid-girdle. There is a
marginal fringe of fine, cylindrical, blunltopped,
spirally grooved spiculés, 112 x 16 p.m. Ven¬
tral scales rectangular, distally rounded, 80 x

Source : MNHN, Paris

MOLLUSCA POLYPLACOPHORA : DEEP-WATER CHITONS

21

Figs 57-67. Ischnochiion (Stenosemus) rohustus sp. nov. : 57. valve IV. dorsal view, x 5. 58. id., rosirai view, x 2.5. —

59. tail valve, dorsal view, x 5. 60. id.. latéral view, x 5. 61, dorsal girdle scale from mid-girdle. latéral view,

x 210. 62. id.. from side margin. ventral view, x 210. 63. id., from mid-girdle, ventral view, x 210. 64.

marginal spiculé, x 210. 65, ventral scales, x 210. 66. central and first latéral radula teeth. x 210. - 67. major

latéral toolh. x 210. 57-60. paratype from Musorstom 4, stn DW 223 (mnhn). 61-67, paralype from Biocal,

stn DW 44 (mnhn).

24 |xm on mid-girdle, shorter towards the outer,
margin, arranged in partly overlapping radial
sériés.

Central tooth of radula rather short, widest in
the proximal half. with a narrow, faintly bilobed
blade ; first laterals relatively longer, with an
excurved blade, projecting exteriorly ; major late¬
rals with a unicuspid dental cap, the cusp curved,
sharply pointed.

Etymology. The Latin adjective rohustus
means solid, relating to the texture of the shell
plates.

Observations. — /. (S.) rohustus differs signifi-
cantly from ail other species in this subgenus by the
ornamentation of the central areas, which some-
what resembles thaï of certain delicately sculpturcd
forms of Chiton tuberculatus Linnaeus, 1758.

Source : MNHN, Paris

22

PI ETE R KAAS

Family SCH1ZOCHITONIDAE
Genus LORICELLA Pilsbry, 1893

Loricella profundior (Dell, 1956)
Figs 68-73

Paricoplax profundior Dell, 1956 : 157, pl. 21, figs 213-
219.

Componochiton raceki Milne, 1963 : 25, figs 1-5.
Loricella oviformis (pars) - K a as. 1985 : 310, figs 41-
45 ; 1990 : 178, figs. Non : Squumophoru oviformis
Nierstrasz, 1905.

Material examined. New Caledonia. Biocal :
stn DW 08, 20"34' S. 166’54' E, 235 m, 12.VIII. 1985 :
1 spm (mnhn).

Musorstom 4 : stn DW 164, 18“33'S, 163“13'E,
255 m, 16.IX.1985 : 1 spm (mnhn). Stn CP 215,
22“56'S, 167“ 17' E, 485-520 m. 28.IX.I985 : 2 spms
(mnhn/1, rmnh/1).

Chalcal 2, stn DW 72, 24“55' S, 168°22' E, 627 m,
28.X.1986 : 1 spm (mnhn). — Stn DW 76, 23'’41'S,
1 67"45' E, 470 m, 30.X.1986 : 4 spms (mnhn/2. rmnh
K2731/1. vb 2987/1). — Stn DW 80. 23°38‘ S,
167"43' E. 435 m, 30.X.1986 : 3 spms (mnhn).

Smib 3 : stn DW 25, 22"56' S, 167“ 16' E, 437 m,
24.V. 1987 : 1 spm (mnhn).

Observations. In 1985 the author estab-
lished the synonymy of the nominal species
Paricoplax profundior Dell, 1956 and Compo¬
nochiton raceki Milne, 1963 with Squamophora
oviformis Nierstrasz, 1905, ranking them with the
genus Loricella Pilsbry, 1893. Now that more
specimens hâve turned up in New Caledonian
waters it becomes obvious that they difler in
several constant features from Nierstrasz's spe¬
cies, which measured 20 x 13 mm, whereas the
Australasian specimens do nol exceed 13 mm in
length. The latéral areas of oviformis show only
7-9 radial ribs, the interstices solid, not perfo-
rated, while there are many more, much finer
riblets, the interstices deeply pitted. in the Aus¬
tralasian and New Caledonian specimens, so that
it appears justifiable to consider them as specifi-
cally different, resulting in the acknowledgement
of Loricella profundior (Dell, 1956) as the valid
name for the Australasian species.

Figs 68-73. — Loricella profundior (Dell, 1965) : 68. head valve, dorsal view, x 10. 69. left hall' of valve II. dorsal view,

x 10. - 70. valve VIII. dorsal view, x 10. 71. id., ventral view, x 10. 72. 73, dorsal girdle scales, x 210. 68-

73. spm from " Vauhan " cruise. 1978/79, sta. 16 (from Kaas, 1985).

Source : MNHN, Paris

MOLLUSCA POLYPLACOPHORA : DEEP-WATER CH1TONS

23

Family CHITONIDAE
Subfamily CHITONINAE

Genus TEGULAPLAX Iredale & Hull, 1926

Tegulaplax pulchra sp. nov.

Figs 74-84

Material examined. — New C'aledonia. Chal-
cal 2 : stn DW 80, 23°27' S, 168°02'E, 160 m,
30.X. 1986 : 1 spm, holotype, now disarticulated
(mnhn).

Diagnosis. — Animal small, 7 x 3.5 mm,
elongate oval, rather highly elevated, carinated.
Valves little beaked, latéral areas well raised,
with 6-8 white, concentric ridges, eut into two
rows of transverse tubercles, one forming the
diagonal ridge, the other accompanying the hind

margin, by a slight radial dépréssion. Central
areas with a narrow, smooth jugal part, ante-
riorly produced, the pleurae with c. 8 deep
longitudinal sulci on either side. End valves with
c. 4-5 white, concentric, undulating ridges, eut
into transverse tubercles by 8-10 vague radial
dépressions. Tegmentum light flesh-coloured,
with roseate blotches and small white dots in the
jugal région. Tail valve with mucro anterior,
antemucronal and postmucronal areas separated
by a tuberculose rib. Apophyses regularly round-
ed, connected across the shallow, convex sinus
by a narrow jugal plate. Girdle roseate banded

Figs 74-84. Tegulaplax pulchra sp. nov. : 74. valve I, dorsal view, x 21. 75. valve IV, dorsal view, x 21. 76. id..

rosirai view, x 21. 77, valve VIII. dorsal view, x 21. 78./</., latéral view, x 21. 79, dorsal girdle scalcs, dorsal

view, x 105. 80./</., ventral view. x 210. 81. marginal spiculé, x 420. 82, ventral scale, x 420. 83, central

and first latéral radula teeth, x 420. 84. head of major latéral tooth, x 420. 74-84. holotype.

Source : MNHN, Paris

24

PIETER KAAS

with white, covered with imbricating, smooth
scales. Dental cap major latéral radula tooth
tricuspid. Gills merobranchial, adanal.

Description. — The unique type measures
6.8 x 3.6 mm, slightly curled before being disar-
ticulated, elongate oval, carinated, dorsal éléva¬
tion c. 0.57, side slopes convex. Head valve semi-
circular, anterior slope straight, the apex slightly
notched, tegmentum ornamented with 4-5 white,
concentric, wavy ridges, eut into transverse
tubercles by c. 8 faint radial dépressions, the
marginal tubercles strongest developed. Interme-
diate valves less than twice as wide as long, the
anterior margin convex, since the central area is
decidedly produced anteriorly ; valve II relatively
longer than the others. Posterior margin almost
straight, the pointed apex barely protruding.
Central area with 6-8 deep, longitudinal, parallel
sulci, with only the innermost not reaching the
front margin. Latéral areas well raised, with up
to 8 white concentric ridges, divided by a slight
radial dépréssion into two rows of transverse,
raised tubercles, the anterior one forming the
diagonal rib, the other one accompanying the
posterior margin. Posterior valve semi-elliptical,
the mucro anterior at about 2/5 the length of the
valve, posterior slope deeply concave, a strongly
developed tuberculose rib separating the antemu-
cronal area from the postmucronal one. Ante-
mucronal area with 5 longitudinal sulci on either
side of the smooth jugum, postmucronal area
with 5-6 wavy, white. concentric ridges.

Articulamentum well developed, white to lighl
roseate, insertion plates narrow. Sût formula

9/1/11, slits inequidistant, slit rays hardly con-
spicuous, teeth finely slriated outside, eaves
finely porous. Apophyses rather short, regularly
rounded, close together, séparated from a nar¬
row jugal plate across the shallow, convex sinus,
by distinct grooves.

Girdle moderately wide, dorsally clothed with
small, imbricating, bluntly rounded, smooth
scales. up to 120 x 80 p.m (mean 108 x 52 [X m),
ventrally with a roundly emarginated base. Mar¬
ginal spiculés smooth, bluntly pointed, 60 x
16 jim, ventral scales elongate rectangular, trun-
cated at both sides, 56 x 12 jxm.

Radula short, c. 1.6 mm long, with c. 25 rows
of mature teeth. Central tooth small, narrow,
bearing an acorn-shaped blade, minor laterals
twice as long, widely diverging, more or less
wing-like, major laterals with a short, tricuspid
head, the cusps bluntly rounded, the central one
longest.

Gills merobranchial, occupying c. 3/4 of the
length of the foot, adanal with interspace, c. 18
ctenidia per side.

Etymology. From the Latin adjective

pulcher = beautiful.

Observations. T. pulchra is the third
known species of this genus. Il is closely related
to T. boucheti Kaas, 1989 from the Philippines,
mainly differing from it in the smooth dorsal
scales (ribbed in boucheti). in the tricuspid head
of the major latéral radula tooth (non-cuspid in
boucheti) and in the anterior position of the
mucro (posterior in boucheti).

Suborder ACANTHOCHITONINA
Family ACANTHOCHITONIDAE
Subfamily ACANTHOCHITONINAE

Genus NOTOPLAX H. Adams. 1861

Subgenus SPONGIOCHITON Dali, 1882

Notoplax (Spongiochiton) produit a
(Carpenter in Pilsbry, 1892)

Spongiochiton productus Carpenter in Pilsbrv, 1892 :

26.

Acanthochites (Notoplax) carpenteri Pilsbry, 1893 : 33
pl. I, figs 14-22.

Acanthochites ( Notoplax) involutus Carpenter in Pilsbry
1893 : 35, pl. I, figs 27-35.

Craspedochiton iiberiensis Thiele, 1909 : 33, pl 4
figs 29-35.

Source : MNHN, Paris

MOLLUSCA POLYPLACOPHORA : DEEP-WATER CHITONS

25

Notoplax foresti Leloup, 1965 : 155, figs 1-3, pis 1-2.
Kaas, 1979 : 873.

Notoplax (Spongiochiton) producia- Kaas, 1989 : 109.

Material examined. New Caledonia. Musor-
stom 4 : stn DW 231, 22°34' S. 167° 10' E. 75 m.
01.X.1985 : 2 spms (mnhn).

Genus CRASPEDOCHITON Shuttleworth. 1853

Craspedochiton hystricosus sp. nov.

Figs 85-96

Material examined. — New Caledonia. Biocal :
stn DW 66, 24°55'S, 168°22' E, 505-515 m. 03.1X.
1985 : 9 spms (MNHN/holotype and 4 paratypes, rmnh
9267/1 paratype, vb 2986/1 paratype, ams/ 1 paratype,
NMNZ/1 paratype).

Chalcal 2 : stn DW 72, 24 n 55' S, 168°22' E. 627 m,
28.X. 1986 : 1 spm. paratype (mnhn). — Stn DW 76,
24°40'S, 168°38' E, 573 m. 29.X.1986 : 1 spm, para-
type (mnhn).

Diagnosis. Animal small, up to 12 x
6 mm, elongate oval, highly elevated. carinated,
valves, including I, sharply beaked, jugal tract
narrow, almost smooth, not distinctly separated
from the evenly granulated latero-pleural areas.
Head valve and latero-pleural areas with only a
vague indication of radial ribs corresponding
with the slits in the insertion plates. Tail valve
small, the insertion plate degenerate posteriorly,
with only a fine irregular dentition. Colour
white. Girdle moderately wide, somewhat en-
croaching at the sutures, narrow posteriorly,
slightly widening anteriorly, finely spiculose.
Suturai tufts small, indistinct. Major latéral
radula tooth with tricuspid dental cap.

Description. Head valve semicircular, an-
terior slope straight to slightly convex, posterior
margin widely V-shaped, with a small. pointed,
protruding apex, outer margin wavy, slightly
curving inwardly between the articulamental
slits, corresponding with them five hardly raised
radiais are to be observed. Tegmentum roughly
granulose, the granules of irregular shape, in-
creasing in size towards the outer margin. ar-
ranged in curved sériés radialing in two direc¬
tions from the apex. Intermediate valves highly
arched, dorsal élévation c. 0.55, strongly cari¬
nated, side slopes straight to a little concave at
both sides of the narrow jugal tract. Front
margin rounded at both sides of the narrow.

concave jugal sinus, side margins very short,
more or less truncated, posterior margin concave
at both sides of the sharply pointed. decidedly
projecting apex. Tail valve small, the length
c. 2/3 of the width. mucro slightly posterior, not
prominent, back slope almost straight, front
margin regularly rounded, antemucronal area
sculptured like latero-pleural areas, postmucro-
nal area like head valve.

Articulamentum thin, white, transparent. In¬
termediate valves with moderately wide, regu¬
larly rounded apophyses, close together, sinus
rather deep, concave, about 1/6 of the width of
the valve, convex in the tail valve, the apophyses
of which are short and trapezoid. Insertion
plates of head valve well developed, with 5 short,
équidistant slits, slit rays hardly perceptible ;
intermediate valves with 1-1 small slits, tail valve
with a very narrow, irregularly and shallowly
toothed posterior insertion plate.

Girdle rather narrow posteriorly, somewhat
widening anteriorly, dorsally coated with fine,
white, straight or slightly bent, bluntly pointed
spiculés, smooth to weakly striated distally, up to
80-100 x 20-25 }j.m. Suturai tufts very small,
composed of c. 40 fine needles, up to 800 x
16 izm. Ventral side of girdle covered with
slightly smaller, straight, weakly longitudinally
ribbed, slender spiculés, 80 x 15 u m.

Radula with a bulging central tooth, bearing
a narrow, straight blade. minor latéral teeth
weakly S-shaped, parallel-sided, partly embrac-
ing the central teeth, major latéral with a
tricuspid blade, the cusps pointed, the central
one much longer than the others.

Gills mcrobranchial, adanal with interspace.

Etymology. The Latin adjective hystrico¬
sus is derived from hystrix ( = porcupine), mean-
ing spinous, relating to the nature of the girdle.

Observations. — C. hystricosus differs from
ail the known Auslralasian and West Pacific

Source : MNHN, Paris

26

PIETER K A AS

species of Craspedochiton by the lack of well tible suturai tufts and especially by its sharply
developed radiating ribs on the head valve and carinated, strongly beaked valves,
latero-pleural areas, by the small, hardly percep-

Figs 85-96. Craspedochiton hystricosus sp. nov. : 85, valve I. dorsal view, x ||. 86. id., ventral view, x H. 87.

intcrmcdiate valve, dorsal view, x il. 88. id., ventral view, x 11.- 89, id., rosirai view, x ||. 90, valve VUE

dorsal view, x 11. 91, id., ventral view, x il. 92, id., latéral view, x ||. 93, head of major latéral radula

tooth. x 230. 94. central and first latéral radula teeth, x 230. 95, dorsal girdle spiculés, x 230. 95, ventral

spiculé, x 230. 85-96, paratype from Biocal, stn DW 66 (mnhn).

Source : MNHN, Paris

MOLLUSCA POLYPLACOPHORA : DEEP-WATER CHITONS

27

REFERENCES

Dall, W, H., 1878. Descriptions of new forms of
mollusks from Alaska contained in the collections
of the National Muséum. Proc. U.S. nain. Mus., 1 :
1-3.

Dell, R. K., 1956. The archibenthal Mollusca of
New Zealand. Dont. Mus. Bull., 18 : 1-235, 265 figs,
2 pis.

Ferreira, A. J., 1985. Chiton (Mollusca : Polypla-
cophora) Fauna of Barbados, West-lndies, with
description of a new species. Bull. mar. Sci.. 36 (1) :
189-219, 18 figs.

Kaas, P., 1979a. On a collection of Polyplacophora
(Mollusca : Amphineura) from the Bay of Biscay.
Bull. Mus. nain. Hist. nat. Paris, (4) 1, sect. A, (1) :
13-31, pis 1-5.

Kaas, P., 1979b. The chitons (Mollusca : Polypla¬
cophora) of Mozambique. Ann. Natal Mus., 23 (3) :
855-879, pis 1-4.

Kaas, P.. 1985. Notes on Loricata (Mollusca) 11-
14. 13, On some little known chitons from the
tropical western Pacific. Zool. Meded. Leiden. 59
(25) : 309-314, figs 41-54.

Kaas, P.. 1989. — Chitons (Mollusca : Polyplaco¬
phora) procured by the Musorstom 3, Philippines
expédition (1985). In : Résultats des Campagnes
Musorstom. Vol. 4 (3). Mém. Mus. nain. Hist. nat.
Paris, (A), 143 : 105-111. figs 6-15.

Kaas, P., 1990. New species and further records of
known species of Polyplacophora from the tropical
Western Pacific. Basteria, 54 : 175-186.

Kaas, P.. & Van Belle, R. A.. 1987. Monograph
of living chitons, 3 : 1-302, 117 figs, 52 maps,
E. J. Brill/Dr. W. Backhuys. Leiden.

Leloup. E.. 1965. — Description d'une espèce nou¬
velle de chiton recueillie par la Calypso dans le
Golfe de Guinée, Notoplax foresti sp. nov. Bull.
Mus. nain. Hist. nat. Paris. (2), 37 (1) : 155-161,
figs 1-3, 2 pis.

Milne, K. L., 1963. A new deep water chiton
(Polyplacophora : Chitonidae) from eastern Austra-
lia. J. malac. Soc. Aust., 7 : 25-27, 5 figs.

Nierstrasz, H. F., 1905. — Die Chitonen der Siboga
Expédition. Siboga Exp., 48 : 1-112, 8 pis.

Pilsbry, H. A.. 1892-1894. Polyplacophora (chi¬
tons). In : G. W. Tryon, Manual of Conchology, 14 :
1-128, pis 1-30 (1892); i-xxxiv, 129-350. pis 31-68
(1893). 15 : 1-64 (1893); 65-133, pis 1-17 (1894).

Richer de Forges, B., 1990. — Les campagnes
d'exploration de la faune bathyale dans la zone
économique de la Nouvelle-Calédonie. In : Résul¬
tats des Campagnes Musorstom. Vol. 6, (1). Mém.
Mus. natn. Hist. nat. Paris. (A). 145 : 9-54, figs 1-13.

Righi, G., 1973. — Adiçôes aos Polyplacophora
brasileiros (Mollusca). Papéis avuls. Zool. S. Paulo,
6 (22) : 259-273, 46 figs.

Thiele, J.. 1909. — Révision des Systems der Chito¬
nen. I. Zoologica Stuttg., 22 : 1-70, figs A-E, pis 1-6.

Source : MNHN, Paris

Source : MNHN, Paris

ILTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSULT

2

Mollusca Bivalvia : Archibenthal
Nuculidae off New Caledonia

Wim BERGMANS

Universiteit van Amsterdam

Instituut voor Taxonomischc Zoologie (Zoôlogisch Muséum)
Postbus 4766, 1009 AT Amsterdam
The Nethcrlands

ABSTRACT

Six spccics of Nuculidae hâve been identified from dredgings
between 250 and 430 m off western New Caledonia : Nucula
nitidulaformis Powell, 1971 : N. kanaka. N. oppressa, and
N. libéra, spp. nov. ; Nucula sp. indet. ; and leionucula

strungei (A. Adams, 1856). Ennucula Ircdalc, 1931, is consi¬
dérai a synonym of Leionucula Quenstedt. 1930. This
nuculid fauna shows rather strong similaritics to that of New
Zealand.

RÉSUMÉ

Mollusca Bivalvia : Nuculidae bathyales de Nouvcllc-Calé-

Six espèces de Nuculidae ont été identifiées dans des
récoltes faites entre 250 et 430 m de profondeur à l'ouest de
la Nouvelle-Calédonie : Nucula nitidulaformis Powell, 1971 ;
N. kanaka. N. oppressa et N. libéra, spp. nov. ; Nucula sp.
indet. ; et Leionucula strungei (A. Adams, 1856). N. nitidula¬
formis et L. strungei ont été décrites de la Nouvelle-Zélande.
N. kanaka montre des similitudes avec N. nitidula A. Adams,
1856. de la Nouvelle-Zélande et des îles Chatham, et avec
N. beachporlensis Vcrco, 1907, de l'Australie. N. oppressa
ressemble à Linucula recens Dell, 1956, de la Nouvelle-

Zélande et des îles Chatham. Seule N. libéra ne se rapproche
pas d'une espèce de la Nouvelle-Zélande et. d'une manière
générale, l'ensemble étudié a des rapports morphologiques
assez étroits avec la faune de Nuculidae de la Nouvelle-
Zélande. Nucula strungei a été classée dans le genre Leionu¬
cula Quenstedt. 1930, qui est caractérisé par l'absence d'une
structure radiale intérieure des valves, qui par suite ont des
bords lisses. Ennucula Iredale, 1931. est considérée comme
un synonyme de Leionucula. Toutes les autres espèces
rapportées possèdent cette structure et le bord ventral crénelé
correspondant, et sont classées dans Nucula Lamarck. 1799.
Dans N. libéra, le bord ventral n’est crénelé qu'à partir d'une
certaine taille.

Bkrgmans, W„ 1991. Mollusca Bivalvia : Archibenthal Nuculidae off New Caledonia. In : A, Crosnihr & P. Bouchet (eds). Résultats
des Campagnes Musorstom. Volume 7. Mèm. Mus. nain. Hist. nul.. (A). 150 : 29-40. Paris ISBN : 2-85653-180-6.

Publié le 20 mars 1991.

Source : MNHN, Paris

30

WIM BERGMANS

INTRODUCTION

On account of my earlier work on a number of
Australian Nuculidae (Bergmans, 1978) Dr Phi¬
lippe Bouchet of the Muséum national d’His-
toire naturelle in Paris invited me to study a
collection of Nuculidae from dredgings from ofT
the west coast of New Caledonia by the vessel
“ Vauban " in 1978 and 1979. As this material

ofiered an obvious opportunity to improve our
knowledge and furlher shape ideas on the distri¬
bution patterns of archibenthal species or species
aggregates in the Australian-Pacific région, and
related taxonomie concepts, I accepted the offer
with gratitude. This paper présents the results of
this study.

MATERIAL AND METHODS

The material on which this report is based is
listed under the species in the taxonomie section.
Descriptions are in accordance with the termino-
logy adopted in Bergmans, 1978, and are ex-
plained in the Method paragraph of that paper.
The présent collection is housed in the Muséum

national d’Histoire naturelle in Paris (mnhn) ;
small samples hâve been deposited in the Zoôlo-
gisch Muséum of the University of Amsterdam
(zma). The photographs in this paper hâve been
made with a scanning électron microscope.

SYSTEMATIC ACCOUNT

Nucula nitidulaformis Powell, 1971
Figs 1-2

Material oxamined. — 1 complété shell. 4 odd
valves, collected by Ph. Bouchet, 23 May 1978, at
" Vauban " Stn 2, 22'17' S, 167°14' E. New C aledonia,
depth 425-430 m (1 shell, 3 valves : mnhn ; 1 valve :
zma).

Description. Valves small, relatively solid,
moderately convex, trapezoid ovate in outline.
Lunule only slightly depressed. Dorsal margin
weakly curved, without médian angulation. Pos-
terior margin truncated, straight ; posterodorsal
angle not very distinct, posteroventral angle
rather distinct. Anterior margin somewhat trun¬
cated, slightly curved ; dorsal margin gradually
passing into anterior margin, anteroventral an¬
gulation broadly rounded but quite distinct.
Ventral margin rounded anteriorly and slightly
less so posteriorly. Exterior of embryonic shell
with probably bifid umbo (shells too worn to be
conclusive) and microscopically pitted surface.
Interdissonconch smooth except growth Unes.

Remaining valve part with some pronounced
growth Unes and on the médian part more or less
aflected by line inner radial structure, which
corresponds with crenulation of ventral margin.
Hinge moderately strong, with up to 8 anterior
and 5 posterior V-shaped secondary teeth (in the
larges! specimens of this sample). Tooth rows
separated by a small chondrophore with a
rounded or slightly angular ventral margin, only
slightly projecling beyond hinge line, directed
downward and very weakly forward. Dorsal
margin somewhat thickened and flattened at
both sides above chondrophore. No discernable
trace of primary teeth. Adductor muscle scars
distinct. Mantle line with shallow incurvation
just anterior of the middle.

Shells white or tinged with pale brown. Traces
of transparent light brown periostracum preser-
ved in two specimens.

Measurements : Table I.

Discussion. Nucula nitidulaformis was de-
scribed from specimens dredged at 366-475 m,

Source : MNHN, Paris

MOLLUSCA BIVALV1A : ARCHIBENTHAL NUCULIDAE

31

Table 1. Measuremcnls in mm and number of sccondary teeth in Nucula niüdulaformis Powell, 1971 from “ Vauban "
Station 2, New Caledonia.

Specimen

Length

Height

Section
(one valve)

embryonic

shell

Number of secondary teeth
anterior posterior

mnhn (right)

1.84

(1.64) '

0.60

8

4

7.MA (left)

1.84

1.60

0.56

8

5

MNHN (right)'

1.76

1.48

7

3

mnhn (doublet)

1.56

1.28

0.44

c. 0.34

MNHN (right)

(1.48)"

1.28

0.44

0.28

6

4

* Ventral margin incomplète ; ' figured spccimen ; " posterior margin damaged.

east of Aldermen Islands, Bay of Plenty, New
Zealand. The smallest specimen in the original
lot, the holotype, measures 4.4 mm in length, the
largest 7.8 mm. Larger specimens appear to be
relatively high. Possible différences between New
Zealand specimens and New Caledonian ones
are the relatively low number of posterior secon-
dary teeth in the latter and the rather perpendi-
cular orientation of its chondrophore. Powell
(1971) counted 10 anterior and 8 posterior teeth
(called posterior and anterior by him, respecti-
vely) in one of the Aldermen Islands specimens

without indicating valve length. He described
the chondrophore as narrow, spoon-shaped, and
anteriorly oblique. His photographs of the holo¬
type specimen suggest that its chondrophore
resembles that of the specimens from New
Caledonia but unfortunately they are not distinct
enough to be conclusive. In this connexion it
may be of importance that Powell did not
assign Nucula niüdulaformis to the genus Pronu-
cula Hedley, 1902 (considered a synonym of
Nucula Lamarck, 1799, by the présent author ;
see Bkrgmans, 1978). Although there is a definite
anteriorly directed vector in its orientation, the
chondrophore in the New Caledonian specimens
possesses the configuration thought to warrant
récognition of Pronucula by Powell. The pré¬
sent sample appears to be the first to be reported
after the description of the species.

Nucula kanaka sp. nov.

Figs 3-4

Material examined. — Holotype : 1 complété shell,
collectcd by Ph. Bouchet, 23 May 1978, at "Vau¬
ban" Stn 2. 22°17'S, 167°14'E, New Caledonia.
depth 425-430 m (mnhn).

Paratypes : 2 left valves, data as for holotype
specimen (mnhn ; zma 3.88.001) ; 1 left valve, collected
by Ph. Bouchet, 23 May 1978, at “ Vauban" Stn 3,
22° 17'S, 167°12'E, New Caledonia. depth 390 m

(MNHN).

Diagnosis. Shell small, triangular. relati¬
vely high, smooth except irregular growth lines.
Chondrophore small but essentially oblique.
Anterior tooth row continuing above chondro¬
phore. Ventral margin crenulated.

Description. — Valves rather small, modera-
tely solid, thickened in the médian area, not
much inflated, rounded triangular in outline.
Lunule not depressed. Dorsal margin weakly
curved, short, gradually passing into long and
nearly straight anterior margin and strongly
descending straight posterior margin. Ventral
margin evenly rounded. Anteroventral angula¬
tion rounded, posteroventral angulation rather
distinct. Umbo pointed. Délimitations of em-
bryonic shell and interdissoconch not discernable
in the présent material. Sculpture consisting of
growth lines and, on the médian area, occasional
radial lines corresponding with a radial inner
structure. Ventral margin crenulated by this
structure. Hinge without a trace of primary teeth
and with rather narrow, V-shaped secondary
teeth. Nine anterior and five posterior teeth in
largest specimen. Anterior tooth row narrowing
towards chondrophore, and continuing with only
one or two very small teeth above it. Chondro¬
phore small, oriented forwards, parallelling an¬
terior tooth row ; its ventral margin slightly
inflated posteriorly and making a rounded angle
with its anterior margin. Adductor muscle scars
shallow but distinct. Mantle line simple. One

Source : MNHN, Paris

Fies 1-8. 1-2 : Nucula nitidulaformis Powcll, length 1.76 mm. from 425-430 m al “ Vauban ” Sln 2 (22"17' S. I67”I4' E),

New Caledonia (mnhn). - .3-4 : Nucula kanaka, sp. nov.. righl valve of hololype specimen, length 2.84 mm. from 425-
430 m at “ Vauban " Stn 2 (22°17'S, 167°I4' E), New Caledonia (mnhn). 5-6 : Nucula oppressa, sp. nov.. lefl valve,
holotype specimen, lcnglh 2.84 mm. from 250-350 m at " Vauban" Sln 40 (22"30‘S. I66"24'E). New Caledonia
(mnhn). — 7-8 : Nucula oppressa, sp. nov.. right valve, paralype specimen, length 2.06 mm. data as for holotype
specimen (mnhn).

Source : MNHN, Paris

MOLLUSCA B1VALVIA : ARCHIBENTHAL NUCULIDAE

33

Table 2. Measuremcnts in mm and numbcr of sccondary teeth in Nucula kanaka, sp. nov., from " Vauban " Station 2
and 3, New Catedonia.

Specimen

Lcngth

Height

Section
(one valve)

Number of sccondary teeth
anterior posterior

Hololype (Stn 2)

2.84

2.48

0.68

right 9
left 8

4

5

Paratype (Stn 2)

2.70

2.44

0.80

9

5

Paratype (Stn 2)

2.40

2.16

0.64

9

4

Paratype (Stn 2)

2.04

1.82

0.60

7

3

Paratype (Stn 3)

2.60

2.25

0.72

8

4

valve with a remnant of a dark brown periostra-
cum.

Measuremcnts : Table 2.

Discussion. — Only the holotype has a well
preserved hinge. In lhe paratypes it is difficult to
see if the chondrophore is rounded and directed
downwards or short and pointing forwards, and
if the anterior teeth ascend above it. Nucula
kanaka resembles the New Zealand species Nucu¬
la nitidula A. Adams, 1856 sensu Powell (1979 :
356, Fig. 83-2), but difiers in outline and details
of the hinge. It is more produced dorsoposte-
riorly, its anterior tooth row is proportionally
much narrower posteriorly, and its chondro¬
phore is much shorter and directed more for¬
wards. Nucula beachportensis Verco, 1907 (see
Bkrgmans, 1978, Fig. 15-19), from Australian
waters, seems also morphologically related to
N. kanaka. N. kanaka difiers from it in being
higher and more triangular, with a proportio¬
nally longer posterior margin, a smaller angle
between its tooth rows, and a shorter chondro¬
phore.

Etymology. The species name kanaka is
derived from “ Kanak ", the original name of the
people of New Caledonia.

Nucula oppressa sp. nov.

Figs 5-8

Material examined. Holotype : 1 left valve,
collected by A. Warén, 7 June 1979. at “ Vauban "
Stn 40, 22°30' S, 166“24' li. New Caledonia. depth 250-
350 m (MNHN).

Paratypes : 54 valves, data as for holotype specimen
(44 valves : mnhn ; 10 valves : zma 3.88.002).

Diagnosis. A small, ovate shell with a
truncated posterior side ; essentially smooth :
with visible inner radial structure in the médian
area and finer, more traverse line structure in
anterior and posterior areas ; typically nuculid
teeth ; a short but very oblique chondrophore ;
and ail margins except the médian part of the
dorsal margin crenulated by the inner structure.

Description. Valves rather small. thin to
moderately solid, médian area thickened in a
number of the specimens — mostly in larger
valves - , moderately inflated, ovate in outline
except for almost perpendicular posterior side.
Radial zone corresponding with straight poste¬
rior margin slightly depressed. Larger valves
relatively higher. Dorsal margin curved. in
some specimens with a slight angle beneath
the umbo. Anterodorsal angulation poorly de-
fined or absent. Posterodorsal angulation round¬
ed but distinct. Anterior margin weakly curved.
partly straight in some specimens. Posterior mar¬
gin short and straight. Anteroventral junclion
weakly marked, rounded, posteroventral junc-
tion more distinct. Umbo pointed. Embryonic
shell bifid, surface finely pitted. Interdissoconch
smooth. Embryonic shell and interdissoconch
without visible radial structure. Remaining part
of valve essentially smooth, with fine growth
fines and in some larger valves a few growth
zones near the margin, mutually separaled by
grooves. Central part of valve with inner radial
structure appearing as fine fines ; anterior and
posterior parts with still finer and widely diver¬
gent fine structure. In a number of specimens.
most of which are quite worn, this finer structure
is hardly or not discernable, especially on lhe
posterior part. Ventral margin crenulated by lhe
radial structure of the central part ; anterior and

Source : MNHN, Paris

34

WIM BERGMANS

posterior margins and adjoining parts of dorsal
margin crenulated by the finer, divaricating
structure. This latter crenulation is difficult to
trace in the majority of the specimens but most
probably distinct in fresh ones. Hinge without a
trace of primary teeth. Dorsal margin slightly
thickened at both sides above chondrophore.
Chondrophore oblique, directed forwards, relati-
vely short. Up to about 12 anterior and five
posterior short, triangular, secondary teeth. An¬
terior tooth row with two or three small teeth
above chondrophore. Adductor muscle scars and
simple mantle line visible in good specimens.
Periostracum not preserved.

Measurements : Table 3.

Discussion. Of the known nuculids, Linu¬
cula recens Dell, 1956, seems to be the morpho-
logically nearest relative of Nucula oppressa.
N. oppressa differs in being less équilatéral, in
having a relatively larger number of anterior
teeth (11 in recens specimens up to 4.04 mm in
length ; Dell, 1956a), in the more oblique and
more elongate form of the chondrophore, and in
the form of the teeth near the chondrophore. In
oppressa , these teeth do not deviate from the
normal nuculid type. In recens “ the two central
teeth of each sériés are larger than their fellows
and meet dorsally under the beaks to give the
appearance of a bifid tooth, the two arms of
which form the latéral limits of the chondro¬
phore ” (Dell, 1956a). (It has occurred to me
that this configuration can only exist in one valve
of each specimen, but its counterpart has not
been described.) Linucula was described as a
subgenus of Nucula by Marwick (1931), to

accommodate small shells from the Tertiary of
New Zealand with weak radial sculpture and
finer radiais on anterior and posterior parts.
Marwick did not describe the hinge, partly
because it had not been preserved well enough in
his specimens but his descriptions were very
short anyway. Dell (1956a) included Nucula
gallinacea Finlay, 1930, in Linucula , on account
of a quite different type of aberrant sculpture on
its anterior side ; his figures of this species show a
hinge typical of Nucula. Dell (1956a) refiected
on the différences between that hinge and the
hinge of Linucula recens , suggesting that Linu¬
cula which he had raised to generic rank
because of the divergent sculpture on its lunule,
its well-defined geographical range, and its long
Tertiary history might consist of two groups.
Bkrgmans (1978) described Nucula brongersmai
from New South Wales, Victoria and Tasmania
in Australia, which also has a sculpture of radial
fines and diverging fines on anterior and poste¬
rior parts, but smooth margins, and a round
chondrophore separating the tooth rows. While
agreeing that forms of aberrant structure and
sculpture of anterior and posterior valve parts
are worth considering in efforts to define natural
groups within the Nuculidae, I do not think that
Linucula is well defined, or that the species with
such structure and sculpture are necessarily
closely related. Further studies of such species
should attempt to describe, and preferably illus-
trate, their hinge configurations in detail.

Etymology. — The spécifie name oppressa ,
derived from the Latin opprimere (to oppress, to
hide) refers to the inner line structures, oppressed

Table 3. Mcasuremcnts in mm and number of secondary teeth in Nucula oppressa, sp. nov., from " Vauban " Station 40.
New Caledonia.

Specimen

Length

Height

(one valve)

Number of secondary teeth
anterior ' ' posterior

Right valve

3.17

2.79

0.92

10

4

Lcft valve '

2.92

2.68

0.87

11

5

Right valve

2.75

2.38

0.75

10

4

Right valve

2.58

2.12

0.71

8

4

Right valve

1.88

1.58

0.50

6

3

Right valve

1.54

1.34

0.42

5

2

Holotypc specimen (Ail others : paratype specimens).
Innermost anterior teeth often worn away.

Source : MNHN, Paris

MOLLUSCA BIVALVIA . ARCHIBF.NTHAL NUCULIDAE

35

in the sense that they do not affect the outer shell
surface (except the margins) and hidden, or
difficult to discern, in many specimens.

Nucula libéra sp. nov.

Figs 9-12

Material examined. Hololype : 1 complété shell.
collected by A. Warén, 7 June 1979. at “ Vauban "
Stn 40. 22°30' S. 166“24' E. New Caledonia. depth 250-
350 m (mnhn).

Para types : 13 complété shells and 246 valves, data
as for holotype specimen (10 shells and 231 valves :
mnhn; 3 shells and 15 valves ; zma 3.88.003).

Diagnosis. — A small, trapezoid ovate shell,
with a relatively long posterior side, rather many
posterior teeth, a partly concave posterior mar-
gin, a smooth outer surface, a weak ovate
chondrophore separating the tooth rows, and in
large specimens a crenulated ventral margin.

Description. Valves small, moderately
solid to solid, thickened in the central area in a
number of larger specimens, only moderately
inflated, trapezoid ovate in outline. Lunule
slightly inflated, demarcated by a radial déprés¬
sion corresponding with a concavity in the
posterior margin. Escutcheon not defined. Dor¬
sal margin straight to slightly concave beneath
the umbo and evenly, although not strongly,
rounded at the sides ; junction with anterior
margin graduai or with a distinct angulation
just above the anteriormost tooth, junction
with posterior margin graduai. Anterior margin
weakly rounded, ventrally straight in some speci¬
mens ; anteroventral angulation rather distinct.
Posterior margin more or less convex along the

tooth row and in most specimens distinctly
concave beneath it ; posteroventral junction dis¬
tinct. Ventral margin rather broadly rounded,
especially posteriorly. Umbo rather pointed.
Embryonic shell extremely small, squarish ovate,
with microscopically pitted surface. Interdisso-
conch smooth, polished, its margin distinct in
part of the specimens. Remaining outer surface
smooth, sometimes, in large specimens, with
some slightly raised growth zones, especially near
ventral margin. Radial structure weakly visible
at a certain angle of light only. Hinge moderately
strong, with up to six anterior and five posterior.
moderately long, secondary teeth. Tooth rows
separated by ovate chondrophore.

Chondrophore wider than high, not much
projecting beyond hinge line, with a weakly
rounded ventral edge. Resilifer roundish triangu-
lar. Dorsal margin thickened at both sides above
chondrophore. in some specimens with traces of
possible vertical striae on these thickened parts.
In large specimens the inner ventral margin is
weakly crenulated. Only in very few specimens a
corresponding structure of radial lines is visible
over a short distance near the crenulations.
Adductor scars visible. Mantle line with shallow
incurvation. Periostracum thin, transparent, tinged
yellowish brown.

Measuremerts : Table 4.

Remarks. — Il is quite unusual that the
crenulation of the ventral margin in an appa-
rently radially structured Nucula shell is présent
only in large, probably adult specimens. and it
is quite confusing when identifying smooth-mar-
gined specimens of this species. The variable
outline, from rather squarish when the antero-
dorsal angle is distinct and the posterior margin

Tabi.f. 4. Measurements in mm and number of secondary teeth in Nucula libéra sp. nov., from " Vauban " Station 40. New
Caledonia.

Specimen

Length

Height

Section
(one valve)

Length
embryonic •
shell

Length

nterdissoconch

Number of secondary teeth
anterior posterior

Paratype. left valve

1.62

1.42

0.38

0.22

4

3

Paralype, right valve

1.84

1.62

0.54

0.22

0.70

7

4

Holotype, right valve

2.18

1.88

0.60

0.22

6

4

Holotype, left valve

6

5

Paratype, right valve

2.21

1.96

0.64

0.20

0.76

7

4

Source : MNHN, Paris

36

WIM BERGMANS

nearly straight, along the tooth row, to trapezoid
ovate when that angulation is not well marked
and the posterior margin distinctly curved, adds
to the identification problem. A further difficulty
is that fresh specimens are transparent and worn
specimens opaque. Moreover, the valves are of
rather variable thickness. It may be problema-
tical to identify individual specimens, and it is
fortunate that the type sériés is so large.

Etymology. This species is named libéra,
meaning free or unrestrained, in memory of Dr
Dick Hillenius, zoologist and writer, for whom
freedom of thought and opinion were synony-
mous with life itself. Dr Hillenius, a colleague
at the Instituut voor Taxonomische Zoologie of
the Universiteit van Amsterdam, died unexpec-
tedly on the 4th of May 1987, at the much too
early âge of 59.

Leionucula strangei (A. Adams, 1856)

Figs 13-14

Nucula strangei A. Adams, 1856 : 52.

Ennucula strangei - Dell, 1956a : 11, pl. 1, fig. I.

Material examined. 31 valves, collected by
A. Warên, 7 June 1979, at “ Vauban " Stn 40,
22°30'S, 166"24' Ei, New C'aledonia. depth 250-350 ni
(26 valves : mnhn ; 5 valves : zma) ; 1 valve, collected
by A. Warên, 7 June 1979, al “ Vauban" Stn 33,
22°33'S, 166"25' H. New Caledonia, depth 290-350 m

(MNHN).

Description. New Caledonian valves up to
about 7 mm long, generally rather thin but
thickened in the area enclosed by adductor
muscle scars and mantle line, moderately in-
flated, broadly trapezoid in outline. Lunule and
escutcheon slightly depressed, lunule bordered by
a weak ridge. Older valves relatively more
convex. Dorsal margin evenly curved, junctions
with anterior and posterior margins graduai.
Anterior margin very slightly convex in small
specimens and straight or with a weak conca-
vity above the junction with the ventral margin
in large specimens. Posterior margin slightly
concave below tooth row. Ventral margin broad¬
ly rounded, posteriorly somewhat flattened, gra-
dually passing into anterior margin but with rath¬
er distinct posteroventral angle. Umbo rather
pointed. Margins of embryonic shell and inter-

dissoconch indistinct in these rather worn speci¬
mens. Embryonic shell microscopically pitted
externally. Rest of valve with fine growth lines,
which in large specimens may become quite
pronounced near the valve margin, where a few
growth zones may develop into concentric
riblets. No trace of an inner radial structure or a
corresponding sculpture.

Anterior hinge plate narrowing towards chon-
drophore and continuing above il. Posterior
hinge plate rather wide, ils inner margin undu-
lated by the teeth. Chondrophore concave, rath¬
er long, projecting beyond hinge plate, oriented
toward anteroventral junction of valve margin ;
in young specimens (Fig. 14) the chondrophore
hardly diverges from the anterior tooth row but
after this stage its orientation changes and il
becomes divergent. Up to 15 anterior and seven
posterior secondary teeth, rather fiat and broad,
with weakly V-shaped basal sections. Adductor
muscle scars distinct. Palliai line distinct, with a
wide curve upward in the middle (Fig. 14). Shells
white or tinged with very pale brown, polished
inside. Periostracum not preserved. Inner ventral
margin smooth.

Measurements : Table 5.

Discussion. Dell (1956b) figured an adult
shallow-water specimen from Wellington Har-
bour, measuring 12.3 mm in length, and remark-
ed that deep-water specimens do not attain a
very large size. The présent specimens are small,
with a number of them about 7 mm in length,
suggesting that this may be the adult size in
the population concerned. C'ompared to Dell’s
figure, they are more trapezoid in outline, with a
somewhat more truncated anterior margin and a
slightly less produced posterior side, and only in
large specimens a chondrophore resembling that
in Dell's specimen. Of the Australian smooth-
margined Nuculidae, Nucula loringii A. Adams
& Angas, 1864 seems related. This species was
synonymized with Nucula cumingii Hinds, 1843
by Hedley (1913), who figured the type speci¬
men, but considered different from that species
by Iredale (1939), who figured a specimen from
Low Isles, Queensland. To judge from those
illustrations, loringii has a more produced poste¬
rior side, a larger angle between the tooth rows,
much weaker teeth, and a narrower and longer
chondrophore. Nevertheless, type material of
loringii and strangei should be compared to

Source : MNHN, Paris

MOLLUSCA BIVALVIA : ARCHIBENTHAL NUCULIDAE

37

Eigs 9-15. — 9-10 : Nucula libéra, sp. nov.. righl valve of paralype specimen. lcngth 2.13 mm. from 250-350 m at “ Vauban "
Sin 40 (22'‘30' S, 166'‘24' E). New Calcdonia (mnhn). 11-12 : Nucula libéra, sp. nov., left valve of paralype specimen.
length 1.71 mm, data as for valve of figs 9-10. 13-14 : Ijeionucula strangei (A. Adams), left valve, length 3.25 mm.

from 250-350 m at " Vauban" Stn 40 (22°30'S. 166"24' E). New Caledonia (mniin). 15 : Nucula sp. indet.. length
2.06 mm, from 425-430 m at “ Vauban" Stn 2 (22° 17’ S, 167° 14' E). New Caledonia (mnhn).

Source : MNHN, Paris

38

W1M BERGMANS

Table 5. Measurements in mm and numbcr of secondary teeth in Leionucula s t rangei (A. Adams, 1856) Irom " Vauban "
Station 40. New Caledonia,

Spécimen

Length

Height

Section
(one valve)

Numbcr of secondary teeth
anterior posterior

Right valve

7.17

5.50

1.83

14

7

Left valve

7.08

5.50

1.83

14

6

Left valve

6.10

4.71

1.25

12

5

Figured left valve

3.25

2.37

0.67

7

3

assess the possible relationship between the two
forms. Unfortunately, Hedley (1913) offered no
discussion on this point.

Nucula sp. indet.

Fig. 15

Material examined. I right valve, collected by
Ph. Bouchet, 23 May 1978. at “ Vauban " Stn 2,
22°17'S, 167 Ü 14'E. New Caledonia. depth 425-430 m

(MNHN).

Description. Valve small, very inequilate-
ral. relatively solid, moderately convex. rounded
triangular in outline. Médian part of valve
thickened. Lunule somewhat depressed, border-
ed by an indistinct radial ridge. Dorsal margin
consisting of an anterior part which is curved at
the ends and straight in the middle, and a very
short, rather straight posterior part ; the two
parts start beneath the umbo at slightly different
levels (the posterior part more ventrally), causing
a slight concavity in the dorsal margin. Antero-
dorsal junction obtuse, posterodorsal junction
more distinct. Anterior margin weakly convex.
Anteroventral junction a broad curve. Posterior
margin weakly convex along tooth row and very
weakly concave below it. Posteroventral angu¬

lation distinct. Ventral margin rounded. Sur¬
face of embryonic shell microscopically pitted.
Outer surface of valve, including interdisso-
conch, smoolh except very fine growth fines.
Hinge rather strong, with seven anterior and
four (fourth minute) posterior, short, secondary
teeth. Anterior tooth row distinctly more narrow
towards chondrophore and continuing above it.
Chondrophore short, not extending far beyond
ventral hinge fine, distinctly forward directed,
with its anterior margin bordering anterior
tooth row but separated from posterior tooth
row by small interspace. Dorsal margin pecu-
liarly thickened along both tooth rows, with
indistinct groove over the length of these, as if
to receive two ridges dorsal to the tooth rows in
the left valve. No trace of primary teeth. Adduc-
tor muscle scars visible. Mantle fine with shallow
incurvation in the middle. Valve white. Traces of
a dark brown periostracum.

Measurements Length 2.06, height 1.84,
section 0.63 mm : length of embryonic shell 0.23
and of interdissoconch 0.56 mm.

Remarks. As there is only one valve of this
apparently undescribed species, and its condition
is not perfect, I prefer to leave it without spécifie
name.

Remarks on the généra Leionucula Quenstedt, 1930, and Ennucula
Iredale, 1931

Iredale’s proposai to place smooth-margined
N ucula species in a separate genus, Ennucula
Iredale, 1931, has been followed by a number of
authors for Australian and New Zealand species.
In his diagnosis of Ennucula, Iredale (1931)
compared it with the type species of Nucula

Lamarck, 1799, i. e. Nucula nucléus (Linnaeus,
1758), only. His diagnosis was very short : “ a
notably oblique chondrophore, above which the
teeth become much smaller, and the angle of
opposition of the two rows of teeth is scarcely
marked ; further, the edge of the European shell

Source : MNHN, Paris

MOLLUSCA BIVALVIA : ARCHIBENTHAL NUCUL1DAE

39

is strongly denticulate, whereas ours is practi-
cally smooth. "

As the obliqueness of lhe chondrophore is
variable in both Nucula and in the species
assigned to Ennucula , and the innermost anterior
teelh, above the chondrophore, are smaller than
their more anterior fellows in ail Nuculidae with
teeth above the chondrophore, the essential
character of Ennucula seems to be its smooth
margin. Quenstedt (1930) had proposed the
genus Leionucula for smooth-margined Nucu¬
lidae. Although that author had not intended a
systematic révision of the Family Nuculidae, his
extensive study of morphological changes in its
représentatives through geological times resulted
in conclusions about some natural sections,
among which Leionucula. The diagnosis of this
section, readily accepted as genus by Thiele
(1934) and later authors, is " Bandgrubenzahn
ôfter, Verbindungsstück der Schlossplatte selten
fehlend. Schalenrand glatt und daher die Grenz-
flâche von Schaloberschicht und Perlmutter
eben. Gault bis jetzt. ” (Quenstedt, 1930 : 112).
(“ Tooth-like projection on hinge plate part in

between resilifer and posterior tooth row often,
hinge plate part in question rarely lacking. Valve
margin smooth and thus interface of outer and
nacreous shell layers smooth. From Gault to
Recent. ”) Diagnostic for ail species is the
absence of radial inner shell structure and the
corresponding smooth margin.

As Iredale offered no additional characters
which would separate Ennucula from Leionucula
— apart from its supposedly different distribu¬
tion in both lime and space (Iredale, 1939) -
1 consider Ennucula Iredale, 1931, a synonym of
Leionucula Quenstedt, 1930. I hâve retained that
genus for Nucula strangei A. Adams for the
simple reason that there is no evidence that the
shells of that species are radially structured,
although it is not altogether impossible that they
are (compare Taylor, Kennedy & Hall, 1969).
Ail other species described in this paper, except
the as yet unnamed single valve, hâve crenulated
ventral margins. Nucula libéra is exceptional in
that this crenulation develops only after a certain
growth stage, but its inner radial structure is
visible from the usual (prodissoconch) stage on.

Affinities of the New Caledonian archibenthal Nuculidae

The Nuculidae treated in this paper represent
lhe first to be reported from New Caledonia.
Two species, Nucula nitidulaformis and Leionu¬
cula strangei , hâve been described from New
Zealand. The latler has been reported from
Australia by Suter (1913) but this has never
been confirmed. In the présent report it is
suggested that strangei should be compared to
the Australian Nucula loringii to assess if and
how closely the two are related.

Nucula kanaka may be related to N. niti-
ilula from New Zealand and Chatham Islands
(Powei.l, 1979) and to N. beachportensis from
southeastem, Southern and northwestern Austra¬
lia (Bergmans, 1978). N. oppressa seems to be
related to Linucula recens from New Zealand
and Chatham Islands (Dell, 1956b). N. libéra is
not obviously related to any of the known New
Zealand species ; it may belong to the same

lineage as the Australian Nucula saltator Iredale,
1939, from Queensland and possibly also from
western Australia (Bergmans, 1978) but that
species is nevertheless quite distinct in a number
of characters (compare Figs 27-29 in Bergmans.
1978). The single valve listed provisionally as
Nucula species cannot be associated with any of
the known species.

It is possible that the New Caledonian archi¬
benthal nuculid fauna, if examined further, will
indeed be found to be more strongly related to
that of New Zealand sensu lato than to others.
At the same time the New Caledonian assem¬
blage may partly consist of endemic species or
species which do occur, or hâve their nearest
relatives, off the Great Barrier Reef. eastern
Papua or Rennell Island, for instance. From
those areas no archibenthal Nuculidae seem to
hâve been described yet.

Source : MNHN, Paris

40

WIM BERGMANS

ACKNOWLEDGEMENTS

I am much indebted to Dr Philippe Bouchet
for his confidence and his sustained patience
during the préparation of this paper. Mr. Rob
G. Moolenbeek of the instituut voor Taxono-
mische Zoologie of the Universiteit van Amster¬

dam made ail the photographs, for which I am
very grateful. He was assisted by the staff of the
Vakgroep Elektronenmicroscopie of the same
university.

REFERENCES

Bergmans. W., 1978. — Taxonomie révision of recent
Australian Nuculidae (Mollusca : Bivalvia) cxcept
Ennucula Iredale, 1931. Rec. Ausi. Mus., 31 (17) :
673-736.

Dell, R. K.. 1956a. Some new off-shore Mollusca
from New Zealand. Rec. Dom. Mus.. 3 (I) : 27-59.
Dell, R. K.. 1956b. The archibenthal Mollusca of
New Zealand. Dom. Mus. Bull.. 18 : 1-235.
Hedley, C., 1913. Studies on Australian Mollusca.
Part XI. Proc. Linn. Soc. N. S. W.. 38 : 258-339,
pis XVI-XDC.

Iredale, T., 1931. — Australian molluscan notes.
No. 1. Rec. Aust. Mus.. 18 (4) : 201-235, pis XXII-
XXV.

Iredale, T., 1939. — Mollusca. Part I. Scient. Rep.

Gt. Barrier Reef Exp.. 5 (6) : 209-245, pis I-VII.
Marwick, J., 1931. — The Tertiary Mollusca of the
Gisborn District. Palaeont. Bull. Wellington. 13:1-
177, pis I-XVII1.

Powell. A. W. B.. 1971. New Zealand molluscan
systematics with descriptions of new species. Part 7.
Rec. Auckland Inst. Mus.. 8 : 209-228.

Powell. A. W. B.. 1979. New Zealand Mollusca.
Marine, land and freshwater shells : i-xiii, 1-500.
Collins, Auckland.

Quenstedt, W.. 1930. Die Anpassung an die
grabende Lebensweise in der Geschichte der Soleno-
myiden und Nuculacecn. Geol. Palaeont. Abh
(n. s.) 18 (1) : 1-120. pis I-Ill.

Suter, H., 1913. Manual of the New Zealand
Mollusca : xxiii, 1-1210. Government Printer, Wel¬
lington.

Taylor, J. D., Kennedy. W. J., & Hall, A., 1969.
The shell structure and mineralogy of the Bivalvia.
Introduction. Nueulacca-Trigonacca. Bu/l. Br. Mus.
nat. Hist., Zool. Suppl.. 3 : 1-125, pis 1-XXIX.
Thiele, J., 1934. Handbuch der systematischen
Weichtierkunde. 3 : 779-1022. Fischer, Jena.

Source : MNHN, Paris

J LT ATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSUL1

3

Mollusca Gastropoda : Seguenziidae
from New Caledonia and the Loyalty Islands

Bruce A. M A RS H A LL

National Muséum of New Zealand
P.O. Box 467, Wellington
New Zealand

ABSTRACT

Thrce subfamilics arc rccognised : Asthelysinae new sub-
family. Seguenziinae Verrill, and Guttulinae Goryachev. Two
tribes are rccognised in Seguenziinae. Fifty five seguenziids
are ncwly rccorded from oIT New Caledonia and the Loyalty
Islands. of which 50 are new to science. These specics are
referable to 13 généra, including 2 new généra and Anxietas

Iredale, which is transferred from Trochidae. Asihelys niti-
dula sp. nov. is bascd on type matcrial from Queensland. Jaw
plates and latcromarginal radular plates are rccorded for the
first time in the family. Seguenziid species richncss and
western Pacific biogeography are briefly discussed.

RÉSUMÉ

Mollusca Gastropoda : Seguenziidae de Nouvelle-Calédonie
et des îles Loyauté.

Une remarquable faune de Seguenziidae est décrite des
étages bathyal et abyssal de la région néo-calédonienne
(bassin des Loyauté et sud de la Nouvelle-Calédonie). La
richesse spécifique de la famille et son apport à la biogéogra¬
phie de l'Ouest Pacifique sont brièvement discutés. L'exis¬
tence de mâchoires et d'une plaque latéromarginale sur la

radula est signalée pour la première fois chez les Seguen¬
ziidae. Cinquante des 55 espèces présentes sont décrites
comme nouvelles de cette région ; une espèce est décrite du
Queensland. Ancistrobasis monodon (Schepman) est transféré
des Calliostomatinac. et Anxietas Iredale des Trochidae, aux
Seguenziidae. Deux nouveaux genres, Eratasthelys et Halys-
tina, et une nouvelle sous-famille, Asthelysinae, sont créés : la
sous-famille nominale Seguenziinae est divisée en deux tribus.

Marshall. B. A.. 1991. - Mollusca Gastropoda : Seguenziidae from New Caledonia and the Loyalty Islands. In : A Crosnier &
P. Bouchet (eds). Résultats des Campagnes Musorsiom. Volume 7. Mém. Mus. nam. Hist. nul.. (A). 150 : 41-109. Paris ISBN : 2-85653-180-6.
Publié le 20 mars 1991.

Source : MNHN, Paris

42

BRUCE A. MARSHALL

INTRODUCTION

During the last ten years the deep-sea gastro-
pods of the family Seguenziidae hâve received
considérable attention from systematists, the
number of généra increasing from 7 to 20, and
the number of Recent species more than dou-
bling to 139 with the présent contribution.
Through studies of shell structure (Bandel,
1979; Barskov, Golovinova & Goryachev,
1980), radula (Marshall, 1983; Quinn, 1983b)
and anatomy (Quinn, 1983b), this formerly
enigmatic family is now firmly established as a
member of the Archaeogastropoda. Salvini-
Pi.awen & Haszprunar (1987) hâve recently
referred the family to a new archaeogastropod
suborder, Seguenziina.

Through the generosity of Philippe Bouchet,
I now hâve the privilège of recording by far
the richest seguenziid fauna known, comprising
55 species in 13 généra. The fact that ail are new
records is testimony not only to the great
diversity of the fauna occurring olf New Caledo-
nia and the Loyalty Islands, but also as to how
little is actually known of the deep-sea molluscan
fauna occurring ofT tropical and subtropical
western Pacific islands in general.

Preserved specimens of several species recorded
living were received too late for inclusion of
descriptions and illustrations of their animais
and radulae.

Abbreviations and text conventions :
ams : Australian Muséum, Sydney;
bmnh : The Natural History Muséum, London;
D : Diameter ;

H : Height ;

mniin : Muséum national d'Histoire naturelle,
Paris ;

nmnz : National Muséum of New Zealand,
Wellington ;

nmp : Natal Muséum, Pietermaritzburg;
nsmt : National Science Muséum, Tokyo ;

TW : Teleoconch whorls (number) ;.
ud : Umbilicus/diameter as percentage of
shell diameter;

usnm : National Muséum of Natural History,
Washington DC ;

zma : Zoôlogisch Muséum, Amsterdam.

Height précédés diameter in ail given dimen¬
sions. Ail shell measuremenls were taken on the
longitudinal axis or at right angles to it. In
descriptions of the posterior notch in the outer
lip, " rétraction deplh ” is the depth from the
adapical insertion to the back of the notch. while
“ prolraction depth ” is the depth from the back
of the notch to the tip of the forward-swinging
abapical part of the lip. Unless otherwise stated
these measurements were taken at the lip rim of
mature specimens.

STATION DATA

Species taken alive are denoted by asterisks.
Station number prefix cp = chalut à perche
(beam trawl), ds = drague type Sanders (epiben-
thic sledge), dw = drague type Waren (rock
dredge).

Biocal campaign stations : N. O. “ Jean-
Charcot

Station DS 04. 21 ° 16' S, 166"40'E, 2 340 m,

11.8.1985 : Asthelys depressa, Halystina cale-
donica *, Seguertzia emmeles.

Station DW 08. 20°34' S, I66°54' E, 435 m,

12.8.1985 : Anxietas exigua, Ancistrobasis
monodon, A. tiara, Calliobasis spectrum, C.
neplicula, C. merista, Fluxinella polita.

Station CP 13. 20"19'S, 167" 18' E, 3 690-

3 740 m, 13.8.1985 : Basilissa superba*.
Station DS 14. 20° 18' S, 167"18'E, 3 680-

3 700 m, 13.8.1985 : Seguenzia platamodes*,
Fluxinella tenera.

Station CP 17. 20"35' S, 167"25'E, 3 680 m,

14.8.1985 : Basilissa superba.

Station CP 23. 22°46' S. I66°20'E, 2 040 m,

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

43

28.8.1985 : Quinnia laetifica*. Seguenzia richeri.

Station CP 26. 22°40' S, 166"27' E, 1618-

1 740 m, 28.8.1985 : Carenzia serrata, C. acan-
thodes, Quinnia limatula, Seguenzia eidalima.
Station DW 33. — 23°10' S, 167*10' E, 675-680 m,

29.8.1985 : Ancistrobasis monodon, Fluxinella
asceta*.

Station DW 36. 23°09' S, 167° 11' E, 650-

680 m, 29.8.1985 : Fluxinella asceta.

Station DW 38. 23*00' S, 167°15'E, 360 m,

30.8.1985 : Calliobasis phimosa, C. spectrum.

Station DW 41. - 22°45' S, 167° 12' E, 380-

410 m, 30.8.1985 : Calliobasis spectrum *.
Station DW 44. 22°47' S, 167*14'E, 440-

450 m, 30.8.1985 : Ancistrobasis tiara*, A.
caledonica*. Calliobasis phimosa*, Fluxinella
poli ta*.

Station DW 46. - 22°53' S, 167°17'E, 570-

610 m, 30.8.1985 : Ancistrobasis tiara*, Fluxi¬
nella polita*, F. asceta*.

Station DW 48. 23°00' S, 167 U 29'E, 775 m,

31.8.1985 : Eratasthelys corona, Ancistrobasis
tiara, A. caledonica, A. adonis. Basilissopsis
charcoti, Fluxinella asceta. F. runcinata.

Station DW 49. 23°03' S, 167°32' E, 825-

830 m, 31.8.1985 : Ancistrobasis adonis*, Fluxi¬
nella runcinata.

Station DW 51. 23°05' S, 167°45' E. 680-

700 m, 31.8.1985 : Ancistrobasis scitula*. Fluxi¬
nella asceta*. F. stirophora.

Station DW 53. — 23°09' S, 167°43'E, 975-

1 005 m, 1.7.1985 : Fluxinella polita, F. asceta,
F. runcinata.

Station DW 56. 23*35' S, 167° 12' E, 695-

705 m, 1.9.1985 : Fluxinella stirophora.
Station CP 57. — 23°44' S, I66°58'E. 1490-
1 620 m, 1.9.1985 : Asthelvs nitidula*. Hadro-
conus grandiosus, Carenzia nitens, C. serrata.
Station DS 59. 23*56'S, 166*’41 ' E, 2 650 m.

2.9.1985 : Asthelys semiplicata, Fluxinella bry-
chia, Basilissa superba, Carenzia ornata, Seguen¬
zia chariessa, S. emmeles, S. levii.

Station DW 64. 24*48' S, 168*09' E, 250 m,

3.9.1985 : Calliobasis festiva.

Station DW 66. 24°55' S, 168*22'E, 505-

515 m, 3.9.1985 : Ancistrobasis monodon.
Station DW 70. - 23*25'S. 167*53'E, 965 m,

4.9.1985 : Fluxinella polita, F. asceta. F. runci¬
nata. Seguenzia eutyches.

Station CP 72. 22*10'S, 167*33'E, 2 100-

2 110 m, 4.9.1985 : Fluxinella brychia, Carenzia
ornata, Quinnia patula, Seguenzia emmeles. S.
levii.

Station CP 75. — 22*19' S. 167*23' E, 825-860 m,

4.9.1985 : Seguenzia chelina, S. chariessa*.

Station DW 77. 22*15'S, 167*15'E. 440 m.

5.9.1985 : Ancistrobasis boucheti*. Fluxinella
membranacea.

Station DW 79. 20*40'S. 166*52'E. 1 320-

1 380 m, 5.9.1985 : Asthelys nitidula, Fluxinella
runcinata, F. euphanes, Carenzia nitens, C.
acanthodes, Halystina carinata, Quinnia lima¬
tula, Seguenzia wareni, S. engonia, S. praeceps,
S. chariessa.

Station DW 80. 20*32'S, 166*48'E, 900-

980 m, 5.9.1985 : Fluxinella asceta, F. runci¬
nata, F. megalomphala, Seguenzia chelina, S.
metivieri, S. matara, S. chariessa. S. stegaslris.
Station DW 83. 20*35'S. 166*54'E. 460 m,

6.9.1985 : Ancistrobasis tiara.

Station DS 98. - 21*24'S, 166*30'E, 2 365-

2 470 m, 7.9.1985 : Carenzia ornata, Quinnia
patula, Halystina caledonica. Seguenzia cha¬
riessa.

Station DW 106. — 21*36'S, 166*29'E, 625-
650 m, 8.9.1985 : Fluxinella xysila.

R. V. “ Vauban ” 1978-79.

Station 40. — 22*30'S, 166*24'E. 250-350 m,
7.6.1979 : Anxietas inspirata, Fluxinella mem¬
branacea, Halystina vaubani, Seguenzia iota.

Source : MNHN, Paris

44

BRUCE A. MARSHALL

SYSTEMATIC ACCOUNT

Subclass PROSOBRANCHIA Milne Edwards, 1848
Order ARCHAEOGASTROPODA Thiele, 1925
Suborder SEGUENZIINA Salvini-Plawen & Haszprunar, 1987
Superfamily SEGUENZIOIDEA Verrill, 1884
Family SEGUENZIIDAE Verrill, 1884

Seguenzidae Verrill, 1884 : 186 (emended). nal anatomy ; three of these groups are here

allocaled subfamilial status, the nominal sub-
The family Seguenziidae is divisible into four family containing two tribes, and are defined as
rather well.defined groups characterised by dis- follows :
tinctive combinations of shell, radula and exter-

Subfamily ASTHF.LYSINAE nov.

Type genus. Asthelys Quinn, 1987.
Included GENERA. Anxietas Iredale, 1917 ;
Thelyssina Marshall, 1983; Asthelys Quinn, 1987;
Eratasthelys gen. nov.

Diagnostic characters. Teleoconch with
anastomosing dendritic threads on first whorl

and/or minute punctations on subséquent whorls.
Axials seldom présent. Posterior notch shallow,
no tooth on inner lip. Snout tip blunt. Central
and latéral teeth stout, rigid, outer marginals
each with fine cusps that extend around lip of
cutting area.

Subfamily SEGUENZIINAE Verrill, 1884

Type genus. — Seguenzia Jefîreys, 1876.

Included généra. - Seguenzia Jeffreys, 1876;
Basilissa Watson, 1879 ; Ancistobasis Dali, 1889 ;
Basilissopsis Dautzenberg & Fischer. 1897 ; The-
lyssa Bayle, 1971 ; Carenziu Quinn, 1983 ; Seguen-
ziopsis Marshall. 1983; Calliobasis Marshall,
1983; Fluxinella Marshall. 1983; Hadroconus
Quinn. 1987; Rotellenzia Quinn. 1987; Quinnia
Marshall. 1988; Halystes Marshall, 1988; Halys-
tina gen. nov.

Diagnostic characters. Teleoconch micro¬
sculpture usually présent, consisting of minute
granulations. Axial sculpture présent or absent.
Posterior notch very shallow to very deep, inner
lip with or without tooth. Snout tip blunt.
Central and latéral teeth stoutly built and rigid,
or thin and flexible, marginal teeth each with fine
cusps that extend around cutting area, or with
very long terminal cusp.

Tribe SEGUENZ1IN1 Verrill, 1884

Included généra. Seguenzia, Carenzia, Diagnostic characters. Posterior notch
Seguenziopsis. Hadroconus, Rotellenzia. Quinnia, of moderate to great depth. Central and latéral
Halystes, Halystina. teeth thin and flexible, marginal teeth each with

very long terminal cusp.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZ1IDAE

45

Table I. Suggcsled phylogcnetic rclationships between seguen/.iid subfamilies, tribes and gênera (not cladistic).

Tribe FLUXINELLINI nov.

Included généra. Basilissa, Ancistrobasis, Central and latéral teeth stoutly built and rigid,

Basilissopsis, Thelyssa, Calliobasis, Fluxinella. outer marginal teeth each with fine cusps that

extend around cutting area.

Diagnostic characters. — Posterior notch
usually shallow, occasionally of moderate depth.

Subfamily GUTTULINAE Goryachev, 1987

Guttulidae Goryachev, 1987 : 23.

Type genus. Guttula Schepman, 1908.

Included généra. — Guttula Schepman, 1908 ;
Sericogyra Marshall. 1988.

Diagnostic characters. Teleoconch entirely
smooth throughout or with microsculpture of

minute granulations. Anastomosing dendritic
threads on first whorl resolving inlo riblets that
are finer and doser than in Asthelysinae and
Seguenziinae. Whorls strongly convex. Apertural
rim simple, withoul notches or tooth. Snout
bifid, tentacular. Central and latéral rather thin
in section, somewhat flexible, marginal teeth
each with long terminal cusp.

REMARKS

Asthelysinae stand well apart from other of wavy dendritic threads on the first teleo-

seguenziids in their distinctive microsculpture conch whorl and/or minute shallow punctations

Source : MNHN, Paris

46

BRUCE A. MARSHALL

that perforate the outer shell layer on sub¬
séquent whorls. Where présent, microsculpture
in other seguenziids consists only of minute
granulations. The significance of these puncta-
tions and granulations is unknown, but they may
facilitate adhesion of the periostracum, or per-
haps they render the shell semipermeable to a
sécrétion by the animal that maintains the
condition of the periostracum. Most Asthely-
sinae are also distinctive in lacking primary axial
sculpture. In the few known axially costate
species the axials gradually résolve after the first
teleoconch whorl, suggesting that axial ribbing
may hâve appeared secondarily in the group, and
thus perhaps independently in Segueziinae. The
radula in Asthelysinae and in the seguenziine
tribe Fluxinellini is distinctive in having stout,
relatively rigid central and latéral teeth, and fine
serrations that extend around the tips of the
outer marginal teeth. By contrast, in the seguen¬
ziine tribe Seguenziini the central and latéral
teeth are thinner in section and flexible, while the
outer marginal teeth each hâve a long, slender
terminal cusp. It seems reasonable to assume
that the radula in Seguenziini has been derived
from the plan exhibited in Asthelysinae and
Fluxinellini through thinning of the central and
latéral teeth, and modification of the marginal
teeth. Although précisé numbers of marginal
teeth per transverse row are difficult to ascertain
accurately from conventional views of the ra¬
dula, the number is certainly greater in Asthely¬
sinae and most Fluxinellini (up to about 20
pairs) than in Seguenziini (up to about 7 pairs),
suggesting that there is a trend toward réduction
in tooth number. If, as here interpreted, Seguen¬
ziini hâve been derived from Fluxinellini, there
would seem to hâve been a trend toward deepe-
ning of the posterior notch and general élabora¬
tion of apertural features throughout the subfa-
mily, which attains peak development in the
genus Seguenzia. Although Seguenziinae are di¬
visible into two groups on the basis of radula
morphology and degree of élaboration of the
apertural features, there is some degree of inter¬
gradation between the groups. For example.

while the shell morphology in Basilissa and
Hadroconus suggests close rclationship to Ancis-
trobasis and Fluxinella (Fluxinellini). the rigidity
of the central and latéral teeth in Basilis.su and
the shape of the marginal teeth in Hadroconus
(Bayer, 1971, fig. 7) are intermediatc between
those in Fluxinellini and Seguenziini. Moreover.
Fluxinella sliropliora and species of Carenzia.
Hadroconus and Quinnia are somewhat interme-
diate in gross shell faciès between elaborately
sculptured Fluxinellini and Seguenzia. Accor-
dingly Fluxinellini and Seguenziini are interpre¬
ted as convenient informai tribal divisions of a
single subfamily in which there is more or less
continuous gradation in shell and radula mor¬
phology. Subfamily Guttulinae is strongly cha-
racterised by the simple shell shape, perfeclly
smooth or distinctively sculptured teleoconch,
and a peculiar snout that is bifid and tentacular
(Marshall, 1988) ralher than blunt-lipped as in
Asthelysinae and Seguenziinae. Guttulinae would
seem to hâve the least derived shell morphology,
with even simpler shells than Asthelysinae. This
simplicity, together with the presence of dendritic
threads in the first teleoconch whorl in Serico-
gyra (Marshall, 1988), suggests a direct relation-
ship with Asthelysinae. The granulate shell micro¬
sculpture on later whorls (Sericogyra) and the
radula, however, are similar to those in Seguen¬
ziini. On the sum of charactcrs and character
States it is concluded that Guttulinae and Seguen¬
ziinae hâve convergent radula morphologies,
that Asthelysinae and Guttulinae diverged early
in the history of the family, and that Seguen¬
ziinae originated from early Asthelysinae (Text
Fig. 1). An equally parsimonious interprétation,
however, is that Seguenziini and Fluxinellini
arose independently from early Guttulinae and
Asthelysinae respectively, which would suggest
that Fluxinellini should perhaps be interpreted as
a subfamily. Since early seguenziids probably
resembled extant Asthelysinae and Guttulinae, it
may be difficult or impossible to unequivocally
recognise fossils among other groups of trochi-
form gastropods.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

47

Subfamily ASTHELYSINAE
Genus ANXIETAS Iredale, 1917

Anxietas Ircdale, 1917 : 334. Type species (by original

désignation) : Anxietas perplexa Iredale, 1917;

Recent, Christmas Island. Indian Océan.

Remarks. Iredale (1917) proposed Anxietas
for a minute barleeid-like gastropod front Chris¬
tmas Island, Indian Océan, the name bestowed
expressing his opinion regarding its relation-
ships. He placed it in Trochidae with considér¬
able réservation. Thiele (1929) considered that
Anxietas was little different from the rissoacean
genus Amphithalamus Carpenter, 1865, Wenz
(1939) considered them synonyms, while Coan
(1964) placed it as a subgenus of Scrobs Watson,
1886 (Barleeidae, Anabathrinae). Ponder (1967)
allowed Anxietas generic rank in Anabathrinae,
but subsequently (1985) reported that A. perplexa
has an internai nacreous layer and returned it to
Trochidae. Although the nacreous layer was not
mentioned by Iredale. its presence may hâve
influenced his decision to place the genus in
Trochidae. Anxietas perplexa is clearly related to
the type species of Thelyssina Marshall, 1983 (T.
sterrha Marshall, 1983), which it resembles in
gross shell morphology, including outer lip profile,
the presence of anastomosing dendritic threads
on the early teleoconch, and minute pits on
subséquent whorls (Marshall, 1983, fig. 5 d, e,
f-h ; Ponder, 1985, fig. 145 a, b). Thelyssina was
referred to Seguenziidae because of similarity to
seguenziids of the généra Ancistrobasis Dali,
1889 and Fluxinella Marshall, 1983 in outer lip
profile. Although Anxietas and Thelyssina are
unknown anatomically, placement in Seguen¬
ziidae is strongly supported by the characteristi-
cally seguenzoid animal and radula in Asthelys
Quinn, 1987 (Marshall, 1988), most species of
which they closely resemble in shell morphology,
including the presence of pits on the teleoconch.
Despite the fact that shell character différences
between Anxietas, Thelyssina and Asthelys are
matters of degree, I prefer to retain the latter two
as distinct généra until animais and radulae can
be compared. They are characterised thus ;
Anxietas — wavy threads on early teleoconch.

no shoulder angulation, suprasutural groove :
Asthelys - no wavy threads or shoulder angula¬
tion on early teleoconch, close, similar peripheral
and suprasutural spiral threads ; Thelyssina —
wavy threads and shoulder angulation on early
spire whorls, strong, rounded peripheral keel.
Note that Anxietas exigua sp. nov. resembles
Anxietas and Asthelys species in lacking a shoul¬
der angulation, yet resembles Thelyssina sterrha
in having a strong, rounded peripheral keel.

Anxietas inspirât a sp. nov.

Figs 1-5, 8

Description. Shell (holotype) 1.85 mm
high, markedly higher than broad, narrowly
trochiform, stout, glossy, anomphalous ; spire
weakly cyrtoconoid, 1.89 x as high as aperture ;
white, nacreous through thin, translucent outer
shell layer.

Protoconch 230 jnn wide, minutely granulate.

Teleoconch of 4.8 whorls, Ist whorl convex,
next whorl becoming fiat, subséquent whorls
very weakly convex, periphery tightly rounded,
base very weakly convex. First whorl sculptured
with fine, crisp, anastomosing. dendritic threads ;
subséquent whorls with fine suprasutural groove
and covered with minute, irregular pits. Base
with 2 fine, close, shallow spiral grooves at about
outer third, absent on last half whorl ; and
strong, rounded, smooth inner spiral cord. Aper¬
ture subtrapezoidal. Outer lip thin at rim, rather
strongly thickened within, posterior notch exlre-
mely shallow and broad, retraction depth 13 %
of shell diameter, almost vertical below apex
before retracting below periphery to shallow,
broad, concave basal notch, no peripheral notch.
Pariétal glaze thin. Inner lip thick, spreading.

Animal unknown.

Type data. — Holotype mnhn (1.85 x
1.40 mm, 4.8 tw) : “ Vauban ", stn 40.

Source : MNHN, Paris

48

BRUCE A. MARSHALL

Figs 1-15. Gênera Anxietas. Asihelys : 1-5, 8, Anxieias inspirata, holotypc, 1.85 x 1.40 mm. 3 x 50. 4 (4lh rw) x 73_
5 x 135,8 x 35. 6, 7, Anxieias exigua, holoiype. 1.60 x 1.65 mm. 9. 10, A. exigua. paralype. Biocal stn DW 08

9 (end Ist tw) x 175. 10 x 120. Il, 12, 14, Asihelys nitidula, holotype, 3.90 x 3.75 mm. 14 (last tw) x s 10 13

15, A. nitidula. Biocal sln DW 79. 3.50 x 3.55 mm. 15 x 90.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

49

Distribution. OIT Southern New Caledo-
nia, 250-350 m (dead).

Remarks. Anxietas inspirata difTers from
(ams) syntypes of A. perplexa in attaining matu-
rity at larger size (1.85 x 1.40 mm. 4.8 tw, cf.

1.60 x |.30. 4.25 tw) and in having a larger
proloconch (diameter 230 |xm, cf. 200 gm). The
two species are otherwise very similar.

Etymology. Inspiring (Latin).

Anxietas exigua sp. nov.
Figs 6. 7, 9, 10: Table 1

Description. Shell up to 2.10 mm high.
glossy, trochiform, rather thin, stout, umbilicus
an elliptical chink, spire 1.36-1.41 x as high as
aperture ; while, nacreous through thin, translu-
cent outer shell layer.

Proloconch 230-240 ;xm wide, minutely granu-
late.

Table 1. Anxietas exigua. Shell measurements (mm) and
countings.

H

D

H/D

TW

2.10

1.70

1.23

4.70

Paratype

1.75

1.03

4.00

Paralypc

1.70

1.00

4.00

Paratype

1.60

1.65

0.97

3.90

Holotype

Teleoconch of up to 4.7 weakly convex whorls.
periphery angulate, base weakly convex. Most of
lst whorl with fine, crisp, anastomosing, dendri-
tic threads, subséquent whorls minutely pitted.
Peripheral keel rounded. adapical edge sharply
shelved and exposed on spire, abapical margin
not defined. Umbilicus bounded by smooth
spiral cord, very narrow, rendered an elliptical
chink by invading inner lip. Aperture subtrape-

zoidal. Outer lip thin at rim, modestly thickened
within ; posterior notch very shallow and broad.
retraction depth 6.9 % of shell diameter, almost
vertical below retracting to broad. shallow.
concave basal notch. No peripheral notch. Pariétal
glaze rather thick. Inner lip curved towards
umbilicus, toothless.

Animal unknown.

Type data. Holotype mnhn : Biocal,

stn DW 08.

Distribution. Off Ouvèa, Loyalty Islands,

435 m (dead).

Remarks. Anxietas exigua difTers from A.
perplexa and A. inspirata primarily in being more
broadly conical and in having a pronounced
peripheral keel. It difTers from Thelyssina sterrha
in being considerably smaller, in lacking a
shoulder angulation on the early teleoconch
whorls and in lacking spiral threads on the outer
part of the base.

Etymology. Small (Latin).

Genus ASTHELYS Quinn, 1987

Asthelys Quinn, 1987 : 66. Type species (by original
désignation) : Basilissa manda Watson, 1879 ; Recent,
eastem Atlantic.

Asthelys nitidula sp. nov.

Figs 11-15, 268-270: Table 2

Description. Shell up to 3.90 mm high,
trochiform, glossy, with narrow umbilical chink,
rather thin, spire 1.19-1.29 x as high as aper¬

ture, white, nacreous through thin, translucent
outer shell layer.

Proloconch 330 um wide, minutely granulate.

Teleoconch of up to 5.20 whorls, lst whorl
strongly convex at first, becoming weakly convex,
subséquent whorls more or less fiat, periphery
angulate, base weakly convex. Sculpture on spire
consisting of 2 close, crisp, smooth, similar spiral
threads, suprasutural spiral commencing on 2nd
half of lst whorl. peripheral spiral partly covered
by succeeding whorls. Fine collabral growth fines

Source : MNHN, Paris

50

BRUCE A. MARSHALL

Table 2. Asthelys nitidula. Shell measurcmcnis (mm) and
countings.

H

D

H/D

TW

3.90

3.75

1.04

5.20

Holotype

3.80

3.55

1.07

5.00

Biocal stn DW 79

3.50

3.55

0.99

5.00

Biocal stn DW 79

3.45

3.65

0.94

4.75

Paratype

and minute circular pits throughout. pits arranged
in spiral lines on Ist few whorls. Base with 2 or
3 similar spiral threads bcside periphery, a
rounded radially pleated spiral cord beside umbi-
lical chink, broad intermediate space with weak
or very weak rounded spiral threads or obscure
spiral lines. Umbilicus very narrow, rendered a
narrow crescenlic chink by invading inner lip.
Aperture subtrapezoidal. Outer lip rim damaged,
thin ; posterior notch very broad and shallow,
retraction depth 5.3 % of shell diameler, slighlly
projected below before retracting to broad, shal¬
low, concave basal notch ; no peripheral notch.
Pariétal glaze thin. Inner lip thick. concave,
spreading into umbilicus, tapered to base, tooth-
less.

Animal unknown (dried).

Radula (Figs 268-270) with the formula c. 12
+ 1 + 1 + c. 12. Central tooth rigid, about as
long as broad, cutting area jutting forward at
right angle from shaft, angulate, with 9-11 sharp,
conical cusps, latérobasal projections prominent.
Latéral teeth rigid, broad, sharply serrate on
inner and outer edges of large, narrowly angulate
cutting area. Marginals slender, each with long
sériés of fine cusps along outer edge of tip and
few on inner edge, inner marginals with strong
terminal cusp, outer marginals with fine cusps
that extend around tip of cutting area. Opercu-
lum thin, multispiral.

Type data. Holotype (ams c. 156439) and
3 paratypes (2 ams, 1 nmnz) : 24°28.2' S,
153°31.2'E, NE of Sandy Cape, Queensland,
Australia. alive. I 330 m-l 380 m. 8 July 1984,
hmas “ Kimbla

Other material examine!) (4 specimens). Bio¬
cal, stn DW 79 (1 mnhn, I nmnz). Stn CP 57
(2 MNHN).

Distribution. Ofif Queensland, Loyalty
Islands. and New Caledonia. I 320-1 620 m, living
at I 330-1 620 m.

Remarks. New Caledonian specimens differ
from the type material in having broader umbili-
cal chinks, but are otherwise indistinguishable on
shell characters. Asthelys nitidula is extremely
similar to A. munda (Watson, 1879) (eastern
Atlantic, 2 058-2 311 m) and A. simplex (Watson,
1879) (off Argentina, 3 475 m) and difTers pri-
marily in having more closely spaced peripheral
spiral cords (see Quinn, 1987, figs 9, 10-14;
Marshall, 1988, figs 1 f-i). It differs further
from A. simplex in having radial pleats beside the
umbilical chink. Compared with A. antaretica
Marshall, 1988 (off South Shetland Islands,
3 715-3 752 m) it differs in having finer periph¬
eral spirals and an almost closed umbilicus with
a radially pleated rim. Asthelys simplex , or a
species very like it, was taken off Westport, New
Zealand at 4 421-4 419 m (Marshall, 1988).

Etymology. Somewhat shining (Latin).

Asthelys semiplicata sp. nov.

Figs 16-20

Description. Shell (holotype) 4.95 mm
high, thin, narrowly trochiform, narrowly umbi-
licate, spire 1.26 x as high as aperture; white,
nacreous through thin, translucent outer shell
layer.

Protoconch 330 p.m wide, surface etched.

Teleoconch of 5.8 whorls; Ist whorl strongly
convex at first, becoming shallowly convex, next
whorl becoming very weakly convex, subséquent
whorls shallowly concave above flattened periph¬
eral keel ; base suddenly contracted, weakly
convex. Peripheral keel comprising 2 close, simi¬
lar, rounded spiral threads, suprasulural spiral
commencing on 2nd half of Ist whorl, peripheral
spiral partly covered by succeeding whorls. Axial
riblets commencing on 2nd whorl, rounded,
collabral, opisthocline, becoming strongly defined
on peripheral keel and rendering it undulant,
weakly defined elsewhere, evanescent on outer
part of base. Minute rounded pits throughout,
arranged in spiral lines on Ist few whorls. Base
with 3 similar spiral threads near periphery, and
3 smooth, rounded spiral cords on inner third,
innermost bordering umbilicus. Umbilicus shallow,
wall tapered. Aperture subtrapezoidal. Outer lip
thin. rim damaged ; posterior notch very broad
and shallow, retraction depth 7.3 % and protection

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

51

16-30, Gcncra Asthelys, Eratasthelys : 16-20, Asthelys semiplicata, holotype, 4.95 x 4.55 mm, 17 (early tw) x 45. 19
(adult periphery) x 45, 20 x 75. 21-25, A. depressa, holotype, 1.40 x 2.22 mm. 24 (base) x 105. 25 x 75 26-Ml

Erarasthelys corona, holotype. 3.65 x 6.70 mm, 29 (end 4th tw) x 35, 30 x 37.

Source : MNHN, Paris

52

BRUCE A. MARSHALL

depth 9.5 % of shell diameter (from collabral
growth lines) ; basal notch broad. shallow, con¬
cave, no peripheral notch. Pariétal glaze thin.
Inner lip thin, gradually tapered to base, tooth-

less.

Animal unknown.

Type data. — Holotype mnhn (4.95 x
4.55 mm, 5.8 tw) : Biocal, stn DS 59.

Distribution. Off Southern New Caledo-
nia, 2 650 m (dead).

Remarks. Asthelys semiplicaia is highly
distinctive among its congeners in having axial
riblets on the teleoconch that are strongly defined
on the peripheral keel. A. semiplicaia resembles
A. muncla in spacing of the spiral threads at the
periphery. but differs from it, and from A.
simplex, in lacking radial pleats at the umbilical
rim. A. semiplicaia further differs from A. sim¬
plex in having a broad, smooth médian band on
the base.

Etymology. — Semiplicate (Latin).

Asthelys depressa sp. nov.

Figs 21-25

Description. - Shell (holotype) 2.22 mm
wide, broader than high, of moderate thickness,
stout, umbilicate, spire 0.71 x as high as aper-
ture ; white, nacreous through thin, translucent
outer shell layer.

Protoconch 330 jj.m wide, finely granulate.

Teleoconch of 3.5 whorls ; subsulural angula¬
tion strong, ramp narrow, more or less horizon¬
tal ; side broad, convex on lst whorl, grading
from fiat to weakly concave on subséquent
whorls ; peripheral keel bluntly angulate ; base
weakly convex. Shoulder angulation commenc-
ing on 2nd half of lst whorl, smooth on lst

2 whorls, strong, rounded nodules on it and
peripheral keel on subséquent whorls ; peripheral
nodules partly covered by succeeding whorls,
their adapical extremities bounded by crisp spiral
thread on last whorl. Summit of peripheral keel
and inner part of umbilical wall minutely granu¬
late, whorls elsewhere pitted, pits arranged in
spiral lines on lst whorl. Base with 2 smooth,
crisp, similar spiral threads beside periphery, and
2 spiral cords beside umbilicus, outer spiral
smooth, innermost strong, with bluntly rounded
nodules, bordering umbilicus. Umbilicus deep,
wall angulate within, outer part shallowly tapered
inwards, inner part steeply tapered outwards,
diameter 26.0 % of shell diameter. Aperture
subtrapezoidal. Outer lip rim damaged, from fine
shallowly sigmoidal collabral growth lines poste-
rior notch extremely broad and shallow, basal
notch shallowly concave, no peripheral notch.
Pariétal glaze thin. Inner lip sharply flexed
toward umbilical wall angulation, rim thin,
rapidly thickened within, very thick against
umbilical wall angulation, evenly tapered to
base, toothless.

Anima! unknown.

Type data. Holotype mnhn (1.40 x
2.22 mm, 3.5 tw) : Biocal, stn DS 04.

Distribution. - Between New Caledonia and
Lifou, Loyalty Islands, 2 340 m (dead).

Remarks. From the teleoconch pitting, A.
depressa appears to be closely related to Anxietas,
Thelyssina, and Asthelys, yet it differs markedly
from members of these généra in the low spire,
wide umbilicus, and nodular spiral cords. A.
depressa is referred to Asthelys because of the
lack of wavy threads on the early teleoconch, but
this placement is obviously provisional.

Etymology. Low (Latin).

Genus ER AT ASTHELYS nov.

Type species. — Eratasthelys corona sp. nov..
Recent, Southern New Caledonia.

Etymology. From the Greek eratos (comely)
and the seguenziid genus Asthelys Quinn, the
latter an anagram of Thelyssa Bayer.

Diagnosis. Shell 6.7 mm wide, sublenticu-
lar, stout, umbilicate, white, nacreous within.
Spire whorls retieulately sculptured with spiral
threads and collabral axial riblets, with addition
of fine, crisp, anastomosing, dendritic threads on
lst 2 teleoconch whorls. Posterior and basal
notches very shallow and broad. Animal unknown.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

53

Remarks. The type species of Eratasthelys
is highly distinctive in combining the dendritic
threads and very shallow labral sinuses charac-
teristic of species of Anxietas and Thelyssina
together with shell shape and reticulate sculpture
similar to those in Ancistrobasis Dali and in
some species of Fluxinella Marshall. On the basis
of similarity in early teleoconch morphology,
Eratasthelys is considered to be most closely
related to Thelyssina. Eratasthelys is probably a
minor olTshoot of the Asthelysinae that has
independently acquired some characteristics of
Ancistrobasis species, rather than a descendant of
the basal stock of Ancistrobasis and related
généra.

Eratasthelys corona sp. nov.

Figs 26-30

Description. — Shell (holotype) 6.70 mm
wide, markedly broader than high, stout, of
moderate thickness. umbilicate ; white, nacreous
through translucent outer shell layer.

Protoconch 330 pm wide, surface abraded.

Teleoconch of 5 whorls, almost fiat above
shallowly concave adapical side of sharply angulate
periphery, last adult whorl becoming weakly and
rather evenly convex, base weakly convex. Axial
riblets weak and indistinct on lst 1.5 whorls,
strong, rounded and fold-like thereafter, con-
fined to adapical half of lst 2 whorls, extending
to periphery on 3rd whorl. Spiral threads more
crisply defined than axials, multiplying by inter¬
calation ; 1 médian spiral surmounting shoulder
angulation on lst 2 whorls, strong at first.

gradually weakening until almost obsolète ; addi-
tional spirals commencing on 3rd whorl, gra¬
dually enlarging. Spirals numbering 8 at end of
2nd to last whorl, including peripheral spiral,
summit of which is partly exposed on spire
whorls. First 2 whorls with fine, crisp, anastomo-
sing, vermiculate threads. Base with 2 similar,
crisply defined spiral cords beside periphery ; fine
inner spiral grooves that become more sharply
defined towards umbilicus : and low, rounded
axial undulations that résolve between periphery
and umbilicus and strengthen towards umbilicus.
Umbilicus deep, wall obscurely spirally lirate.
diameter 26 % of shell diameter. Aperture sub-
trapezoidal. Outer lip of moderate thickness.
posterior notch very broad, extremely shallow.
retraction depth 4.03 % and protraction depth
1.98 % of shell diameter ; basal notch broad and
shallow, no peripheral notch. Pariétal glaze thin.
Inner lip thick, rim tightly folded towards umbi¬
licus, concave below insertion, almost straight
below, toothless.

Animal unknown.

Type data. Holotype mnhn (3.65 *
6.70 mm, 5 tw) : Bioc al, stn DW 48.

Distribution. — Off Southern New Caledo-
nia, 775 m (dead).

Remarks. An extremely distinctive species
combining shell features of Thelyssina and Ancis¬
trobasis species (see above).

Etymology. — Crown (Latin).

Subfamily SEGUENZIINAE
Tribe FLUXINELLINI

Genus ANCISTROBASIS Dali. 1889

Ancistrobasis Dali, 1889 : 383. Type species (by
subséquent désignation of Dall, 1927) : Basilissa
costulata Watson, 1879; Recent, south-eastern Flo¬
rida and Gulf of Mexico.

Ancistrobasis monodon
(Schepman, 1908) comb. nov.

Figs 31-35

Calliostoma (Astele) monodon Schepman, 1908 : 68,
pl. 6, fig. 2.

Type data. - Figured syntype zma 3.08.082
(4.70 x 5.75 mm, 5.25 tw) : “ Siboga ", stn 95,
5°43.5' N, 119°40' E, off Sabah. Malaysia. 522 m.

Other material examined (3 specimens mnhn). —
Biocal, stn DW 08 (1). Stn DW 33 (1).

Stn DW 66 (1).

Distribution. OIT Sabah. Malaysia, and
soulhem New Caledonia. 505-680 m (dead).

Source : MNHN, Paris

54

BRUCE A. MARSHALL

Figs 31-45. Genus Ancistrobasis : 31-35, Ancislrobasis monodon. Biocai. stn 66. 5.70 x 7.90 mm, 34 (last TW) x 20,
35 x 70. 36-40. A. liara, hololype. 2.70 x 3.31 mm. 39 (last rw) x 35. 40 x 100. 41-45, A. calédonien, hololype,

4.05 x 5.05 mm, 44 (last tw) x 25, 45 x 80.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

55

Remarks. — New Caledonian specimens are
indistinguishable from the holotype of Callios-
toma (Asiele) monodon a typical Ancistrobasis
species in sculpture, shape, and shell size
relative to the numbcr of whorls. The présent
specimens difier, however, in having slightly
larger proloconchs than the holotype (diameter
330 [j.m, cf. 310 ij.m), and in attaining maturity at
larger shell size (5.70 mm x 7.90 mm. 5.9 tw),
and may prove to represent a distinct, closely
relaled species. See Discussion page 107.

Ancistrobasis tiara sp. nov.

Figs 36-40; Table 3

Description. Shell up to 3.55 mm wide,
broader than high, stout, of moderate thickness,
umbilicate, spire 1.41-1.59 x as high as aper-
ture ; white, nacreous through thin, translucent
outer shell layer.

Protoconch 270-280 um wide, ralher coarsely
granulate.

Teleoconch of up to 5.25 whorls. Shoulder
angulation strong on lst whorl. weakening and
becoming obsolète on 2nd whorl, strongly supra-
median at first, descending to submedian posi¬
tion ; subséquent whorls weakly convex ; base
suddenly contraeted below angulate, gently undu-
lant periphery, weakly convex. Spire whorls
sculptured with prominent, rounded, shallowly
sigmoidal, collabral axial riblets, almost obsolète
on peripheral spiral, interspaces about twice as
wide as each axial, traversed by numerous finer
spiral threads thaï mulliply by intercalation.
Submedian spiral and one submedially between
it and periphery strong, others finer and similar,
interspaces smooth apart from fine collabral
growth fines. Base sculptured with low, rounded
collabral riblets that extend to umbilical rim. and
9-11 spiral cords; outer 2 or 3 spirals narrow.

Table 3. Ancistrobasis tiara. Shell measurements (mm)
and countings. (Biocal, stn DW 44).

Character

„

Range

Mean

SD

H

10

2.65-2.91

2.75

0.08

D

10

3.13-3.55

3.35

0.12

U D

10

0.80-0.86

0.82

0,02

TW

10

5.00-5.25

5.07

0.10

UD%

10

22.6-28.5

25.0

1.66

most prominent, distinctly nodular. outer 3 or

4 spirals with interspaces considerably wider
than each spiral ; inner spirals doser, broadening
towards umbilicus with narrowing interspaces,
innermost spiral very broad, with rounded radial
pleats. Umbilicus deep, diameter 22.6-28.5 % of
adult shell diameter. Aperture subquadrate. Outer
lip thin at rim of labial projection, elsewhere
thick ; posterior notch broad. retraction depth
4.98-5.55 % and projection depth 6.95-9.25 % of
shell diameter ; basal notch concave, no periph¬
eral notch. Pariétal glaze thin. Inner lip thick.
slightly tapered at abapical extremity. toothless.

Animal unknown (dried).

Type data. Holotype (2.70 x 3.31 mm.

5 tw) mnhn, and 20 paratypes (ams, bmnh.
MNHN. NMNZ, NMP, USNM) : BlOCAL, Stn DW 44.

Other material examined (13 specimens mnhn).
Biocal, stn DW 08 (5). Stn DW 46 (5).

Stn DW 48 (1). Stn DW 83 (2).

Distribution. Off Ouvéa. Loyalty Islands,
and Southern New Caledonia. 435-775 m, living
at 440-610 m.

Remarks. Ancistrobasis tiara closely resem-
bles the Kermadec species A. dilecta Marshall.
1983, from which it diflfers primarily in having a
considerably stronger shoulder angulation on the
early teleoconch whorls, while the secondary
spirals are considerably more numerous. It dilTers
from A. monodon in being smaller, in having a
shallow posterior notch. in having a shallowly
instead of deeply undulant peripheral keel, and
in lacking a denticle at the inner base of the
outer lip. A. tiara and A. monodon occurred
together at Biocal stn DW 08.

Etymology. Crown (Latin).

Ancistrobasis calcdonica sp. nov.

Figs 41-45; Table 4

Description. Shell up to 5.90 mm wide.
broader than high, stout, of moderate thickness.
umbilicate, spire 1.70-2.00 x as high as aper¬
ture ; white. nacreous through translucent outer
shell layer.

Protoconch 330 um wide. coarsely granulate.

Source : MNHN, Paris

56

BRUCE A. MARSHALL

Teleoconch of up to 5.8 whorls. lst 2 whorls
with distinct shoulder angulation, strong on
lst whorl, weakening and becoming obsolète on
2nd whorl, angulation descending from strongly
supramedian to a submedian position, sub¬
séquent whorls weakly convex, periphery angu-
late, rendered strongly undulant by axial riblets,
base weakly convex. Spire whorls sculptured
with prominent, rounded, shallowly sigmoidal,
collabral axial riblets that extend to periphery,
interspaces slightly wider than each axial, traversed
by finer spiral threads that multiply by intercala¬
tion, 9-11 major spirals at start of last adult
whorl, small, rounded nodules al intersections,
bases of interstitial pits finely granulate on later
whorls. Base sculptured with low, rounded colla¬
bral riblets that extend to umbilical rim, and 10
or 11 similar, rounded spiral cords ; interspaces
on outer part about twice as wide as each spiral,
narrowing towards umbilicus, intersections with
low, rounded nodules, innermost spiral more
strongly beaded. Umbilicus deep, diameter 22.6-
26.8 % of adult shell diameter. Aperture sub-
quadrale. Outer lip rather thin at rim. thickened
within, a spiral thickening near base of inner lip
forming a prominent, rounded denticle at rim ;
posterior notch very broad and shallow, rétrac¬
tion depth 6.78-7.98 % of shell diameter, descen¬
ding more or less vertically before retracting to
shallow, concave basal notch.

Animal unknown (dried).

Table 4. Ancistrobasis calédonien. Shell mcasurements (mm)
and countings. (Biocal, stn DW 44).

Character

n

Range

Mean

SD

H

10

3.90-4.45

4.23

0.20

D

10

5.05-5.90

5.46

0.27

H/D

10

0.73-0.82

0.77

0.03

TW

10

5.50-5.80

5.62

0.11

UD%

10

22.6-26.8

24.8

1.34

Type data. Holotype (4.05 x 5.05 mm,
5.5 tw) mnhn, and 27 paratypes (ams, bmnh,
MNHN. NMNZ, NMP, L'SNM) : BlOCAL, Stn DW 44.

Other material examined (2 specimens mnhn).
Biocal. stn DW 48.

Distribution. OfT Southern New Caledo-
nia, 440-775 m (alive).

Remarks. Ancistrobasis caledonica difTers
from the sympatric A. tiara sp. nov. and the
Kermadec A. dilecta Marshall in having a larger
protoconch, stronger spirals and doser axials on
the teleoconch, and a more strongly undulant
periphery, while the intersections are more dis-
tinctly nodular. Il difTers further from A. tiara in
having a denticle at the inner base of the adult
outer lip. From A. monodon, which it resembles
in size, shape and in having a denticle at the
inner base of the outer lip, A. caledonica difTers
in having broader, more closely spaced axial
riblets, stronger nodules, and finer interstitial
granules on the spire. A. caledonica and A.
monodon hâve overlapping depth ranges oIT
Southern New Caledonia and the two species are
probably locally sympatric.

Etymology. (New) Caledonian.

Ancistrobasis scitula sp. nov.

Figs 51-55, 271 : Table 5

Description. Shell up to 6.10 mm wide,

broader than high, stout, of moderate thickness,
umbilicate, spire 1.43-1.46 x as high as aper¬
ture : white, nacreous through translucent outer
shell layer.

Protoconch 350-370 gm wide, finely granulate.

Teleoconch of up to 5.80 whorls, lst 3 whorls
with supramedian shoulder angulation, subséquent
whorls almost fiat ; periphery narrowly rounded,
rendered strongly undulant by axial riblets ; base
weakly convex. Spire whorls sculptured with
prominenl. rounded, sigmoidal, collabral axial
riblets, interspaces at least twice as wide as each
axial, traversed by numerous spiral threads that
multiply by intercalation, major spirals num-
bering about 7 at start of last adult whorl, a
subsutural spiral vanishes at end of lst whorl.
Small. rounded. conical nodules at intersections,
finely granulate at bases of interstitial depres-

Tabi.e 5. — Ancistrobasis scitula. Shell measurcments (mm)
and countings.

H

D

H/D

TW

UD%

4.70

5.53

0.85

5.80

19.9

Paralypc

4.50

5.50

0.82

5.75

25.4

Holotype

4.35

5.25

0.83

5.60

20.9

Paratype

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

57

Figs 46-60. - Genus Ancisirobasis : 46-50, Ancistrohasis boucheti. holotype, 3.10 x 5.71 mm. 49 (last tw) x 125. 50 x 70. —
51-55, A. scitula, holotype, 4.50 x 5.50mm. 54 (last tw) x 25. 55 x 70. 56, 57, 59, 60, A. adonis, holotype.

3.20 x 6.50 mm, 59 (last tw) x 30, 60 x 90. 58, A. adonis, paratype. Biocal stn DW 49, width 6.10 mm.

Source : MNHN, Paris

58

BRUCE A. MARSHALL

sions on later whorls. Shoulder spiral and spiral
near abapical quarter strongest and similar on
lst 4 whorls, after which secondary spirals
become as strong as shoulder spiral, that near
abapical quarter remaining stronger. Base sculp-
tured with narrow, rounded collabral riblets that
are evanescent immediately within umbilical rim,
and 9-11 stronger, rounded spiral cords, outer
3 with interspaces considerably wider than each
spiral, interspaces narrowing towards umbilicus
until about as wide as each spiral, small rounded
nodules at intersections, innermost spiral strongly
beaded at umbilical rim. Umbilicus deep. rim
angulate, diameter, 19.9-25.4% of adult shell
diameter. Aperture subquadrate. Outer lip thin
at rim, thickened within posterior notch, little
thickened elsewhere ; posterior notch broad, shal-
low, retraction depth 8.2 % and prolraction
depth 5.2. % of shell diameter ; basal notch
broadly concave, shallow, no peripheral notch.
Pariétal glaze thin. Inner lip thick, simple, gently
tapered to abapical extremity.

Animal. Snout little over twice as long as
broad, rounded latéral projections at tip, mouth
a vertical sût beneath. Cephalic tentacles beside
snout, inner bases at least twice as far apart as
width of each tentacle base, narrowly tapered,
dorsoventrally flattened, edges ciliate, left slightly
longer than snout. right yet slightly longer, large
black eyes at outer bases. Right suboptic tentacle
slightly larger than right cephalic tentacle, longi-
tudinally grooved beneath, the groove directly
above tall, thin longitudinal ridge. Epipodial
tentacles large, 6 on each side.

Radula (Fig. 271). Central looth rigid, about
as long as broad, cutting area jutting forward at
right angle from shaft, angulate, with 7-9 stout,
conical cusps, médian cusp largest ; latérobasal
projections prominent. Latéral teeth stout, broad,
cutting area of each angulate, cusps sharp,
terminal cusp largest, 5 or 6 smaller cusps on
outer edge and 2 or 3 on inner edge. Marginal
teeth slender ; innermost tooth broader than
outer teeth, tip narrowly angulate, sharply ser-
rate ; outer marginals each with finely and
narrowly serrate. angulate tips, and long sériés of
sharp slender cusps along outer edge.

Jaw plates thin, subrectangular, broader than
long, éléments short, anteriorly elongating.

Type data. Holotype mniin and 5 para-
types (4 mniin. 1 nmnz) : Biocal, stn DW 51.

Distribution. OIT Southern New Caledo-
nia, 680-700 m (alive).

Remarks. Ancistrobasis scitula resembles A.
tiara in lacking a tooth at the inner base of the
outer lip at maturity, and diflers in being more
strongly sculptured with a more strongly rounded
periphery. It difiers from ail other Ancistrobasis
species in details of teleoconch sculpture, in its
exceptionally large protoconch, and in having a
subsulural spiral thread on the first teleoconch
whorl. To my knowledge, jaw plates hâve not
hitherlo been recorded from this family. They
are recorded herein from Fluxinella, Calliobasis
and Basilissa. They are also présent in at least
one species of Seguenzia (S. compta Marshall,
1983).

Etymology. Beautiful (Latin).

Ancistrobasis houcheti sp. nov.

Figs 46-50, 272-277 ; Table 6

Description. Shell up to 6.03 mm wide,
considerably broader than high, very stout,
rather thick, umbilicate, spire 1.00-1.13 x as
high as aperture ; white. nacreous through trans¬
itent outer shell layer.

Protoconch 330-370 ^m wide. surface granu-
late.

Tablk 6. Ancistrobasis houcheti. Shell mcasuremcnls (mm)
and countings. (Biocal. sln DW 77).

Charactcr

n

Range

Mcan

SD

H

10

2.50-3.20

2.96

0.19

D

K)

4.45-6.03

5.57

0.43

H/D

II)

0.48-0.56

0.53

0.02

TW

10

4.40-5.10

4.84

0.21

UD%

10

24.7-29.3

27.3

1.55

Teleoco

nch

of up to 5.10

whorls, lst

whorl

with strong, strongly supramedian angulation,
angulation weakening over next hall' whorl and
vanishing while descending to médian position,
subséquent whorls weakly concave ; periphery
sharply angulate, rendered sharply and closely
serrate by axial riblets ; base weakly convex.
Spire whorls sculptured with prominent, roun¬
ded, flexuous collabral axial riblets that exlend

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZ1IDAE

59

to periphery, summits broad, flattened. inter-
spaces about 1.5 x as wide as each axial, tra-
versed by numerous, considerably finer spiral
threads, of which peripheral spiral is strongest,
others similar ; intersections not nodular, enlire
surface covered with minute granules after Ist
whorl. Base sculptured with rounded. flexuous
collabral riblets that extend to umbilical rim, and
10 or 11 rounded spiral cords, interspaces wider
than each spiral, outer 3 spirals similar. slightly
narrower and doser than others, which are
similar to each other, small, rounded nodules at
intersections ; innermost spiral strongly beaded at
umbilical rim. Umbilicus deep, rim angulate,
diameter 24.7-29.3 % of adult shell diame-
ter. Aperture subquadrate. Outer lip thick at
rim, strongly thickened within, with prominent,
angulate elliptical ridge against posterior notch,
strong outer basal ridge, and prominent, rounded
elliptical denticle near base of inner lip. inter¬
spaces concave, deeply so in front of labial
projection. Posterior notch shallow, broad, rétrac¬
tion depth 3.50-4.60 % and protraction deptli
4.72-5.84 % of shell diameter ; basal notch broad
and shallow. no peripheral notch. Pariétal glaze
rather thick. Inner lip very thick. suddenly
tapered at base to form bluntly rounded denticle.

Animal. Snout considerably longer than broad,
sides subparallel, broad, rounded latéral exten¬
sions at tip, mouth a vertical slit below. Cephalic
tentacles parallel to snout, lying above it, slightly
longer than snout, right tentacle slightly longer
than left, inner bases almost in contact, tapered,
dorsoventrally flattened, edges and ventral sur¬
face ciliate, large black eyes at outer bases. Right
suboptic tentacle larger than cephalic tentacles,
dorsoventrally flattened, ventral surface longitu-
dinally grooved directly above a narrow tentacle
that emerges from its base. Epipodial tentacles
large, tapered, 8 right and 7 left. Opercular lobe
small, operculum thin. chitinous, spiral.

Radula (Figs 272-274) with the formula
c. 13 + 1 +1 + 1 +c. 13. Central tooth rigid,
about as long as broad, cutting area jutting
forward at right angle from shaft, angulate, with
9 large, sharp, narrowly conical cusps, médian
cusp largest. Latéral teeth stout, broad, cutting
area of each angulate and sharply serrate. Inner¬
most marginal long and narrow, stout, cutting
area narrowly angulate, terminal cusp very large,
subterminal cusps numbering about 7 on outer
edge, fewer on inner edge. Outer marginals

slender, each with numerous sharp slender cusps
at tip and along outer distal edge.

Jaw plates (Figs 275-277) subrectangular.
broader than long, thin, éléments enlarging
anteriorly, their tips minutely and sharply rough-
ened.

Type data. — Holotype (3.10 x 5.7 1 mm.
5.10 tw) mnhn and 17 paratypes (ams. bmnh,
NMNZ, NM P, USNM) : BlOCAl-, stn DW 77.

Distribution. — Off Southern New Caledo-
nia. 440 m (living).

Remarks. Ancistrobasis boucheti is extre-
mely distinctive in the combination of low spire,
strongly thickened shell, shallowly concave spire
whorls, and flattened axial riblets crossed by
numerous much finer spiral threads.

Etymology. I take particular pleasure in
naming this superb species after Philippe Bou¬
chet.

Ancistrobasis adonis sp. nov.

Figs 56-60 ; Table 7

Description. Shell up to 6.65 mm wide,
considerably broader than high, stout, of mode-
rate thickness, umbilicate, spire about 1.25 x as
high as aperture ; white. nacreous through trans¬
itent outer shell layer.

Protoconch 320-330 jxm wide, surface slightly
roughened, almost smooth.

Table 7. Ancistiobasis adonis. Shell measurements (mm)
and countings. (Biocai., stn DW 49).

Character

n

Range

Mean

SD

H

10

2.45-3.65

3.10

0.33

D

10

4.90-6.65

6.06

0.59

H/D

10

0.49-0.55

0.51

0.02

TW

10

4.70-5.60

5.20

0.23

UD%

10

26.8-33.8

30.2

2.16

Teleoconch of up to 5.60 whorls ; periphery
angulate, rendered shallowly serrate by axial
riblets : base weakly convex. Shoulder angulation
strong on lsl whorl, descending from strongly
supramedian to submedian position, weak and at
about abapical third on subséquent whorls.

Source : MNHN, Paris

60

BRUCE A. MARSHALL

becoming obsolète on last adult whorl. Spire
whorls almost fiat after Ist whorl, last adult
whorl weakly convex, a low, immediately subsu-
tural angulation develops late on 2nd whorl and
becomes finely beaded, becoming obsolète on
last adult whorl. Spire whorls sculptured with
fine, crisp, sigmoidal, collabral axial riblets, these
traverser! by fine, crisp spiral threads that mul-
tiply by intercalation, numbering 16-18 at start
of last adult whorl, thread surmounting shoulder
angulation strongest, others finer and similar,
interspaces finely granulate on ail whorls. Basal
axials flexuous, rounded, collabral, vanishing on
outer part of umbilical wall, becoming almost
obsolète on last adult whorl. Basal spirals
numbering 14-18 in adults, interspaces finely
granulate, outer 4 or 5 narrow and prominent,
about as strong as axials, widely spaced, inner
spirals lower than axials inwardly progressively
widening then narrowing, interspaces narrower
than each spiral ; innermost 2 spirals narrowest,
widely spaced. connected by rounded radial
pleats that extend onto outer part of umbilical
wall. Umbilicus deep, rim angulate, diameter
26.8-33.8 % of adult shell diameter. Aperture
subquadrate. Outer lip thin at rim, slightly

thickened within ; posterior notch shallow, broad,
retraction depth 3.03-3.07 % and protraction
depth 4.54-4.61 % of shell diameter ; basal notch
concave, no peripheral notch. Pariétal glaze thin.
Inner lip thick, simple, gently tapered abapically,
toothless.

Animal unknown (dried).

Type data. Holotype (3.20 * 6.50 mm,
5.3 tw) mnhn and 51 paratypes (ams, bmnh,
MNHN, NMNZ, NMP, USNM) : BlOCAL, Stn DW 49.
Paratype (1 mnhn) : Biocal, stn DW 48.

Distribution. OfT Southern New Caledo-
nia, 775-830 m, living at 825-830 m.

Remarks. Ancistrobasis adonis is rendered
highly distinctive by ils low spire, fine axial
riblets, internally slightly thickened outer lip,
persistent shoulder angulation, subsutural angu¬
lation. and by the immédiate appearance of
secondary spirals and minute granules on the
teleoconch.

Etymology. After Adonis, a beautiful
youth beloved by Venus.

Genus BASILISSOPSIS Dautzenberg & Fischer, 1897

Basitissopsis Dautzenberg & Fischer, 1897 : 163. Type
species (by monotypy) : Basitissopsis watsoni Daut¬
zenberg & Fischer, 1897 ; Recent, northeastern
Atlantic.

Remarks. — The new species described below
and Ancistrobasis regina Marshall, 1983 closely
resemble B. watsoni in general faciès and thus
appear to be closely related. Their shells are
essentially similar lo those of Ancistrobasis spe¬
cies, and differ primarily in having a strong
shoulder angulation on ail teleoconch whorls.
Ancistrobasis species hâve a shoulder angulation
on the earliest teleoconch whorls that soon
becomes obsolète, so this différence is clearly a
matter of degree. Basitissopsis may eventually
prove to be better treated as a subgenus of
Ancistrobasis , or perhaps a synonym, but I prefer
to maintain it at generic level until animais and
radulae can be compared. An undescribed spe¬
cies occurs in Otaian (Early Miocene) beds at
Parengarenga Harbour, northern New Zealand.

Basitissopsis charcoti sp. nov.

Figs 61, 62, 64-65

Description. Shell (immature holotype)
1.60 mm wide, broader than high, thin. umbili-
cate, spire 1.18 x as high as aperture, white,
nacreous through thin, translucent outer shell
layer.

Protoconch 320 (im wide, smooth.

Teleoconch of 3 strongly shouldered whorls,
shoulder angulation strongly supramcdian at
first, descending to an almost médian position,
ramp almost fiat ; side steeply tapered, weakly
concave : periphery sharply angulate ; base weakly
convex. Spire whorls axially and spirally orna-
mented. Axial riblets rounded, widely spaced,
flexuous. collabral, interspaces with very fine
granules and faint collabral growth fines, entirely
traversing whorls, weak at periphery, prominent
elsewhere. Shoulder and peripheral spirals strong,
about as strong as axials, shoulder spiral with

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

61

Figs 61-75. Gênera Basilissopsis. Calliobasis : 61, 62, 64, 65, Basilissopsis eharcoti, hololypc, 1.12 x 1.60 mm. 64 (last
TW) x 100, 65 x 90. — 63, 66-70, Calliobasis phimosa, 2.50 x 5.25 mm, 63 x 25, 69 (last tw) x 50. 70 x |() 5 . 7 |-

75, C. festiva, holotype, 2.52 x 2.85 mm, 74 (last tw) x 45 , 75 x 105.

Source : MNHN, Paris

62

BRUCE A. MARSHALL

small, rounded conical nodules where inter-
secting axials, peripheral spiral gently undulant,
summit exposed on spire on lst 2 whorls then
covered by succeeding whorl ; interspace with
2 fine spiral threads that traverse axials. Base
with 10 spiral cords, outer 3 narrow and widely
spaced, inner spirals broader, closer and tra-
versing weak, rounded, axial riblels. Umbilicus
deep, rim rather sharply angulate, diameter 29 %
of shell diameter. Aperture subquadrale. Outer
lip thin, simple, posterior notch (from collabral
sculpture) very broad and shallow, very slightly
retracted from suture, weakly protractive below ;
basal notch broad and concave. Pariétal glaze
very thin. Inner lip thin, straight, simple.

Animal unknown.

Type data. Holotype mnhn (1.12 x

1.60 mm, 3 tw) : Biocal, stn DW 48.

Distribution. Off Southern New Caledo-

nia, 775 m (dead).

Rkmarks. Among nominate species, Basi-
lissopsis charcoti most closely resembles B. regina
(Marshall, 1983) from oit the Three Rings
Islands, northern New Zealand (622-805 m),
dilTering primarily in lacking spiral threads on
the ramp. The Atlantic B. watsoni has a much
stronger peripheral keel. A species very similar to
B. charcoti. perhaps the same, is represenled by
two fragmentary specimens from the northern
Three Rings Risc (nzoi stn U602, 31"30.7'S,
172'’49.8'E, 1216-1 385 m). Judging from the
simple aperlural features. and particularly the
size of the protoconch, the holotype of B.
charcoti is immature, as are probably ail of the
specimens of Basilissopsis species known from
northern New Zealand.

Etymology. After Biocal campaign ship
N. O. “ Jean-Charcot ”.

Genus CALLIOBASIS Marshall, 1983

Calliobasis Marshall, 1983 : 254. Type species (by

original désignation) : Basilissa bombax Cotton &

Godfrey, 1938; Recent, Southern Australia.

Remarks. — Calliobasis and Ancistrobasis
species are similar in gross shell and external
animal morphology. The radula of C. spectrum
sp. nov. (see below) differs from that in Ancistro¬
basis and ail other known seguenziids in having
lateromarginal plates probably reduced inner
marginal teeth and in having a shallowly
rounded rather than angulate cutting area on the
latéral teeth with relatively much larger cusps.
Calliobasis species differ further from Ancistroba¬
sis in attaining smaller shell size and in being
smaller relative to the number of whorls (maxi¬
mum diameter 2.25-3.01 mm. 4.80-5.50 teleo-
conch whorls, cf. 3.55-7.90 mm. 5.10-5.90 teleo-
conch whorls). Moreover, they hâve fewer inter-
calating spiral threads on the spire, and stronger
peripheral and suprasutural spirals, while the
shoulder angulation tends to be more persistent.
Apart from C. nepticula sp. nov., the available
specimens of which are possibly bleached, the
species are outstanding among known seguen¬
ziids in having yellowish green or greenish yellow
shell pigmentation, which suggests an unusual

diet. Unlike typical Ancistrobasis species, intersti-
tial granulation commences immediately after
the protoconch. The gap in shell morphology
between Calliobasis and Ancistrobasis species is
bridged to some extent by Ancistrobasis adonis
sp. nov., and A. scinda sp. nov., the former
having interstitial granules on ail teleoconch
whorls, A. scitula having both a distinct suprasu¬
tural angulation and a persistent shoulder angu¬
lation. Although Calliobasis and Ancistrobasis
are undoubtedly closely related, I prefer to
maintain Calliobasis as a genus rather than a
subgenus of Ancistrobasis , primarily because of
the distinctive radular morphology and because
both groups hâve been separate since at least the
Eocene ( Calliobasis eos Marshall. 1983 and
Ancistrobasis pacifica Ladd, 1970).

Calliobasis phimosa sp. nov.

Figs 63, 66-70; Table 8

Description. Shell up to 2.80 mm wide,
about as broad as high ; spire broadly conical'
1.17-1.60 x as high aperture. stout, of moderato
thickness, umbilicus invaded by inner lip.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

63

Colour of protoconch and Ist 1.5-1.75 teleo-
conch whorls pale lime green. Subséquent whorls
white, nacreous through thin. translucent outer
shell layer. narrowly axially maculated at about
each 4th or 5th axial costa with greenish yellow,
maculations occasionally darkening to yellowish
brown on last adult whorl, pigmentation deepest
on spirals ; base white.

Table 8. — Calliohasis phimosa. Shell measuremcnts (mm)
and countings. (Biocal. stn DW 44).

Charactcr

n

Range

Mcan

SD

H

10

2.15-2.60

2.45

0.12

D

10

2.40-2.80

2.63

0.13

H/D

10

0.87-1.01

0.93

0.04

TW

10

5.00-5.50

5.24

0.19

UD%

10

20.3-24.0

22.4

1.19

Protoconch 270 ^m wide. coarsely granulate.

Teleoconch of up to 5.5 convex whorls, sub-
sutural ramp weakly convex, base suddenly
contracted, very weakly convex. Spire whorls
sculptured with strong. similar, reticulating spiral
cords and collabral axial costae, interspaces
concave, sharp conical nodules at intersections,
minute granules throughout. Spiral cords num-
bering 3 or 4 on last adult whorl. Shoulder spiral
commencing immediately, at about adapical third ;
suprasutural spiral commencing early 2nd whorl,
rapidly enlarging to resemble shoulder spiral :
intermediate spiral (présent in about 1 specimen
in 3, including holotype) commencing on last
adult whorl. remaining weaker than others ;
peripheral spiral covered by succeeding whorls,
weaker than shoulder and intermediate spirals.
Base sculptured with collabral axial riblets thaï
extend into umbilicus, and 8 or 9 spiral cords,
outer 2 spirals narrowesl, outer 3 widely spaced.
innermost spiral strongest, beaded at umbilical
rim, low rounded nodules at intersections with
axials. Umbilicus shallow, conical. infilled by
inner lip, diameter 20.3-24.0 % of adult shell
diameter. Aperture subquadrate. Outer lip thin
at rim of labial projections elsewhere strongly
thickened, especially behind peripheral and basal
notches ; posterior notch gently flared, retraction
depth 4.87-6.07 % and protraction depth 10.98-
13.34% of shell diameter. Forward-swinging
limb depressed adaperturally, basal notch smal-
ler than anterior, rim slightly flared ; peripheral
notch very small, very shallowly retracted. Parié¬

tal glaze thin. Inner lip thick. spreading into
umbilicus.

Animal unknown (dried).

Type data. — Holotype mnhn (2.50 x
5.25 mm, 5.25 tw) and 25 paratypes (ams, bmnh.
mnhn, nmnz, nmp. usnm) : Biocai., stn DW 44.
Paratypes (3 mnhn) : Biocal, stn DW 38.

Distribution. — Off Southern New Caledo-
nia, 360-450 m, living at 440-450 m.

Remarks. Calliobasis phimosa dilfers from
hitherto named species of Calliohasis in having
an infilled umbilicus, and in sculptural details,
particularly in the late appearance or absence of
the intermediate teleoconch spiral.

Etymology. From the Greek phimos (stop-
ping an orifice) and refering to the infilled
umbilicus.

Calliohasis festiva sp. nov.

Figs. 71-75

Description. — Shell (holotype) 2.85 mm
wide. slightly broader than high ; spire broadly
conical, 1.5 x higher than aperture, deeply umbili-
cate, stout, of moderate thickness.

Protoconch translucent white. First 1.5 teleo¬
conch whorls reddish brown through translucent
outer shell layer, next whorl translucent white.
Succeeding whorls translucent, regularly axially
maculated with yellowish brown, each 3rd or
4th nodule on peripheral spiral deeply pigmen-
ted. coinciding nodule on shoulder spiral more
lightly pigmented, very lightly pigmented in a
spiral band between shoulder spiral and médian
spiral. Base translucent white. Aperture nacreous
within.

Protoconch 260 gm wide, coarsely granulate.

Teleoconch of 5.25 whorls. rather strongly
convex at first, becoming weakly convex, sub-
sutural ramp weakly convex, base suddenly
contracted, very weakly convex. Spire whorls
sculptured with strong, similar, reticulating spiral
cords and collabral axial costae, interspaces
concave, sharp conical nodules at intersections,
minute granules throughout. Spiral cords num-
bering 7 on last adult whorl. Shoulder spiral
commencing immediately, at about adapical quar-

Source : MNHN, Paris

64

BRUCE A. MARSHALL

ter, suprasutural spiral commencing on 2nd half
of lst whorl, rapidly enlarging to resemble
shoulder spiral ; intermediate spiral commencing
late on 3rd whorl, remaining weaker than shoul-
der and suprasutural spirals ; ramp spiral com¬
mencing late on 4th whorl. becoming about as
strong as intermediate spiral ; peripheral spiral
slightly weaker than suprasutural spiral, covered
by succeeding whorls ; 2 spirals intercalate near
end of first half of last adult whorl, one between
shoulder and intermediate spiral, the other bet¬
ween intermediate spiral and suprasutural spiral.
Base sculptured with collabral axial riblets that
extend into umbilicus, and 8 spiral cords, low
rounded nodules at intersections, innermost spi¬
ral beaded at umbilical rim. Umbilicus conical,
deep, diameter 25 % of adult shell diameter.
Aperture subquadrate. Outer lip strongly thick-
ened within ; retraction depth of posterior notch
7.02 % of shell diameter, protraction depth
unknown (labial projection broken) ; basal notch
concave, peripheral notch very small. Pariétal
glaze thin. Inner lip thick, gently curved, sud-
denly tapered near abapical extremity to form a
small projection.

Animal unknown.

Type data. Holotype mnhn (2.52 x

2.85 mm, 5.25 tw) : Bioc al, stn DW 64.

Distribution. Off Southern New Caledo-

nia, 250 m (dead).

Remarks. — Calliobasis festiva seems closest
to the Kermadec C. miranda Marshall, 1983,
from which it differs in attaining larger size, in
the presence of a spiral cord on the ramp, and in
having a stronger shoulder spiral and doser
suprasutural and peripheral spirals. Il differs
from C. phimosa sp. nov. in having an open
umbilicus and in sculptural details, notably in
the presence of a spiral cord on the ramp, and in
the early appearance of the intermediate spiral.

Etymoi.ogy. Delightful (Latin).

Calliobasis spectrum sp. nov.

Figs 76-80, 278, 279 ; Table 9

Description. Shell up to 2.26 mm wide,
slightly broader than high, stout, openly umbili-
cate, spire 1.26-1.56 x as high as aperture.

Protoconch and lst 2.75 whorls white, sub¬
séquent spire whorls white, narrowly axially
maculated with yellow at about each 4th or 5th
axial costa, base white, aperture nacreous within.

Protoconch 230-260 (j.m wide, coarsely granu-
late.

Tabli: 9. Calliobasis spectrum. Shell mcasurements (mm)
and countings.

H

D

H/D

TW

UD%

2.02

2.13

0.95

4.80

23.4 Biocal stn

i DW 41

2.00

2.26

0.88

5.00

28.0 Biocal stn

i DW 08

1.91

2.23

0.86

4.90

25.4 Holotype

1.90

2.20

0.86

4.75

30.3 Biocal stn

i DW 08

Teleoconch of up to 4.9 whorls, convex at
first, becoming weakly convex, subsutural ramp
more or less fiat, base suddenly contracted,
weakly convex. Spire whorls sculptured with
strong, similar, reticulating spiral cords and
collabral axial costae, interspaces concave, rounded
conical nodules at intersections, minute granules
throughout. Spiral cords numbering 6 on last
adult whorl. Shoulder spiral commencing imme-
diately, at about adapical quarter, relatively
weak ; suprasutural spiral commencing on lst half
of lst whorl, larger than shoulder spiral after
lst whorl ; intermediate spiral commencing late
on lst half of 4th whorl, enlarging to resemble
shoulder spiral ; adapical ramp spiral commencing
on 3rd whorl, abapical ramp spiral commencing
of 2nd half of 4lh whorl, both enlarging to
resemblc shoulder spiral : peripheral spiral covered
by succeeding whorls, intermediate in size be¬
tween shoulder and intermediate spirals. Base
sculptured with collabral axial riblets that extend
into umbilicus, and 7-9 spiral cords, low rounded
nodules at intersections, innermost spiral beaded
at umbilical rim. Umbilicus conical, deep, diame¬
ter 23.4-30.3 % of adult shell diameter. Aperture
subquadrate. Outer lip thin at rim of labial
projection, elsewhere strongly thickened ; poste¬
rior notch shallow, retraction depth 6.24-8.96 %
of shell diameter : basal notch concave, no
peripheral notch. Pariétal glaze thin. Inner lip
thick, tapered at abapical extremity to form a
small projection.

Animal white. Snout about twice as long as
broad, tip deeply cleft between strong, rounded
latéral projections, mouth a vertical slit below.
Cephalic tentacles similar, dorsoventrally flat-

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

65

Figs 76-90. Gcnus Calliobasis : 76-80, Calliobasis spectrum, holotype, 1.91 *2 21 mm 79 (last rw)
î* 5 ’ < ' J' e P ,icu,a - holotype, 2.30 x 3.10 mm. 84 (lasl tw) x 45. 85 x 95.' 86-90.

2.10 x 2.40 mm, 89 (last tw) x 45. 90 x 100.

x 60.80 x 105. 81-

C. merista, holotype.

Source : MNHN, Paris

66

BRUCE A. MARSHALL

tened, ciliate, gradually tapered, inner bases
about 2 tentacle base-widths apart, large swollen
black eyes at outer bases. Right suboptic tentacle
as large as cephalic tentacles, posteroventral
surface concave. Epipodial tentacles large, 6 on
each side. Operculum thin, chitinous, spiral.

Radula (Fig. 278). Central tooth rigid, about
as long as broad, tip flattened, cutting area
jutting forward at right angle from shaft, angu-
late, with about 9 conical cusps, médian cusp
largest, latérobasal projections prominent. Laté¬
ral teeth stout, broad, each with broad, shallowly
rounded cutting area with 9 or 10 relatively
large, long, narrow cusps. Innermost marginal
reduced to form convoluted articulatory latero-
marginal plate. Outer marginals slender, small,
sharp, slender cusps at tips and in long sériés on
outer edges.

Jaw plates (Fig. 279) thin, ovate, broader than
long, éléments short, longer anteriorly.

Type data. — Holotype and 2 paratypes
mnhn : Biocal, stn DW 41. Paratypes (3) :
Biocal, stn DW 08 (1 mnhn, 1 nmnz) ; Biocal,
stn DW 38 (1 mnhn).

Distribution. Off Ouvéa, Loyalty Islands
and Southern New Caledonia, 360-435 m, living
at 380-410 m.

Remarks. Calliobasis spectrum superficially
resembles the South Australian C. bombax
(Cotton & Godfrey, 1938) (Marshall. 1983, fig.
7 g), from which it differs primarily in attaining
maturity at considerably smaller size, and in the
later appearance of spiral cords on the spire
other than the shoulder and suprasutural spirals.
From the New Caledonian C. fesliva sp. nov.,
which also has a fully open umbilicus, it differs in
having weaker spiral cords above the suprasutu¬
ral spiral, which is set higher on late whorls, and
in the later appearance of the intcrmediate spiral.
It differs from the locally sympatric C. phimosa
sp. nov. in having a fully open umbilicus and in
sculptural details.

Etymology. Image (Latin).

Calliobasis nepticula sp. nov.

Figs 81-85

Description. - Shell up to 3.01 mm wide,
broader than high, stout, openly umbilicate.

spire rather broadly conical, 1.30-1.35 x as high
as aperture. White (bleached ?), aperture nacreous
within.

Protoconch 270 «m wide, coarsely granulate.

Teleoconch of up to 4.8 whorls, spire whorls
convex at first, becoming very weakly convex :
suturai ramp fiat throughout or becoming shal¬
lowly concave : base suddenly contracted, weakly
convex. Spire whorls sculptured with crisp, reti-
culating spiral cords and arcuate, collabra! axial
costae, interspaces concave, small conical nodules
at intersections, minute granules throughout.
Spiral cords numbering 6 on last adult whorl.
Shoulder spiral commencing immediately, at
about adapical quarter; suprasutural spiral com¬
mencing almost immediately, becoming slightly
larger than shoulder spiral ; intermediate spiral
commencing on 2nd half of 3rd whorl, enlarging
to resemble shoulder spiral ; subsutural spiral
commencing at start of 2nd half of 3rd whorl,
remaining weaker than shoulder spiral ; spiral
between shoulder and intermediate spirals com¬
mencing on 2nd half of 4th whorl, enlarging
to resemble adjacent spirals ; peripheral spiral
covered by succeeding whorls, similar to supra¬
sutural spiral. Base sculptured with arcuate
collabral axial riblets thaï are evanescent on
outer part of umbilical wall, and 7-9 similar,
widely spaced spiral cords, low rounded nodules
at intersections ; innermost spiral radially pleated
at umbilical rim. Umbilicus conical, deep, diame-
ter 25.4-31.0 % of adult shell diameter. Aperture
subquadrate. Outer lip thin at rim of labial
projection, elsewhere strongly thickened ; poste-
rior notch broad, retraction depth 4.4 % and
protraction depth 5.0 % of shell diameter ; basal
notch concave, no peripheral notch. Pariétal
glaze thin. Inner lip rather thin, a small rounded
denticle near base.

Animal unknown.

Type data. Holotype mnhn (2.30 x
3.10 mm, 4.8 tw), and 2 paratypes mnhn, nmnz
(2.40 (est.) x 3.00 mm, tw ? ; 2.10 x 2.87 mm,
4.6 tw) : Biocal, stn DW 08.

Distribution. Off Ouvéa, Loyalty Islands,
435 m (dead).

Remarks. Calliobasis nepticula closely
resembles C. bombax in general faciès, but
differs in attaining maturity at smaller size, in

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZ11DAE

67

being more broadly conical, and in having more
densely crowded granules on the spire whorls. It
differs from C. phimosa sp. nov. in having a fully
open umbilicus and in sculptural details, from C.
fesliva in having a weaker shoulder spiral and
more widely spaced suprasutural and peripheral
spirals, and from C. spectrum in being more
finely sculptured, and in having one spiral cord
on the subsutural ramp. The lack of shell
pigmentation is possibly a distinctive character,
but the available specimens are obviously long
dead and so may well be bleached.

Etymology. Young granddaughter (Latin).

Calliobasis mais ta sp. nov.

Figs 86-90; Table 10

Description. Shell up to 2.40 mm wide,
about as broad as high ; spire 1.33-1.79 x as
high as aperture, stout, openly umbilicate.

Colour of protoconch and lst 2 teleoconch
whorls pale greenish yellow. Subséquent whorls
white, narrowly maculated at each 4th or 5th axial
costa with greenish yellow, most deeply pigmented
on suprasutural spiral, base white. aperture
nacreous within.

Table 10. — Calliobasis meristu. Shell measurcments (mm)
and countings. (Biocal, sln DW 08).

Character

«

Range

Mcan

SD

H

6

2.05-2.25

2.15

0.07

D

6

2.25-2.40

2.34

0.05

H/D

6

0.87-0.%

0.92

0.04

TW

6

4.75-5.25

5.03

0.21

UD%

6

21.2-33.3

27.3

3.93

Protoconch 270 u.m wide, coarsely granulate.

Teleoconch of up to 5.25 whorls, markedly
convex at first, becoming weakly convex ; ramp
narrow, weakly convex or fiat ; base suddenly
contracted, very weakly convex. Spire whorls
sculptured with strong. similar. reticulating spiral
cords and sigmoidal, collabral axial costae, inter-

spaces concave, conical nodules at intersections,
minute granules throughout. Spiral cords num-
bering 4 on last adult whorl. Shoulder spiral
commencing immediately, at about adapical quar-
ter ; suprasutural spiral commencing on 2nd half
of lst whorl, rapidly enlarging to resemble
shoulder spiral ; intermediate spiral commencing
on 2nd half of 4th whorl, remaining weaker than
adjacent spirals ; peripheral spiral similar to
suprasutural spiral, summit covered by suc-
ceeding whorls. Base sculptured with collabral
axial riblets that extend into umbilicus, and
8 spiral cords, outermost close beside peripheral
spiral, interspaces of outermost 4 spirals widest,
other interspaces about as wide as each spiral,
innermost spiral beaded at umbilical rim, other
spirals with low rounded nodules at intersections
with axials. Umbilicus conical, deep. diameter
22.2-33.3 % of adult shell diameter. Aperture
subquadrate. Outer lip thin at rim of labial
projection, elsewhere strongly thickened, poste-
rior notch broad, retraction depth 5.50-5.80 %,
and protraction depth 8.00-9.00 % of shell dia¬
meter, basal notch concave, no peripheral notch,
distinctly notched against umbilical rim. Pariétal
glaze thin. Inner lip thick, tapered at abapical
extremity, toothless.

Animal unknown.

Type data. Holotype mnhn (2.10 x
2.40 mm. 4.90 tw) and 7 paratypes (1 ams.
5 MNHN, 1 NMNZ) : BlOCAl., stn DW 08.

Distribution. Off Ouvéa, Loyalty Islands,
435 m (dead).

Remarks. Calliobasis merista is most simi¬
lar to C. phimosa in shape and sculpture,
differing primarily in being openly umbilicate, in
having stronger and sharper nodules on the
shoulder and suprasutural spirals, and in having
the shoulder spiral set lower on the adult whorls.
C. merista occurred with C. spectrum and C.
nepticula at the type locality.

Etymology. — Divided (Greek).

Source : MNHN, Paris

68

BRUCE A. MARSHALL

Genus FLUXINELLA Marshall, 1983

Fluxinella Okutani. 1968 : 42 (nomett nudum).
Fluxinella Marshall, 1983 : 250. Type species (by
original désignation) : Fluxinella lepida Marshall.
1983 ; Recent, New Zealand.

Fluxinella membranacea sp. nov.

Figs 91, 93, 94, 97; Table 11

Description. Shell up to 4.00 mm wide,
considerably broader than high, sublenticular.
thin, fragile, glossy, umbilicate, spire 0.78-1.00 x
as high as aperture ; white, nacreous through
thin, translucent outer shell layer.

Protoconch 280-300 um wide, side very finely
granulate, summit essentially smooth.

Table II. Fluxinella membranacea. Shell measure-
ments (mm) and countings. (" Vaubun". stn 40).

Charactcr

n

Range

Mcan

SD

II

10

1.22-1.75

1.49

0.16

D

10

3.20-4.00

3.58

0.23

Il D

10

0.38-0.47

0.41

0.03

TW

10

3.90-4.50

4.31

0.26

UD%

10

27.0-30.3

28.8

1.15

Teleoconch of up to 4.5 whorls. Shoulder
angulation sharp on Ist whorl, progressively
weakening until obsolète near end of Ist half of
3rd whorl. summit level with adapical extremity
of protoconch on Ist half whorl, gradually
descending over subséquent whorls to suprame-
dian position. Second and later spire whorls
weakly convex, shallowly concave above and
below sharp, very shallowly undulant peripheral
keel. base well rounded. Spire smooth apart from
fine collabral growth fines. Base smooth apart
from collabral growth fines and obscure spiral
fines. Umbilicus deep, rim narrowly rounded,
projecting inwards to overhang almost vertical
wall. diameter 27.0-30.3 % of adult shell diame-
ter. Aperture subrhomboidal. Outer lip thin and
fragile, damaged in ail available specimens, pos-
terior notch very shallow and broad, from
growth fines weakly retracted from suture and
weakly projected below ; basal notch concave ;

peripheral notch contained in keel, not retracted.
Pariétal glaze very thin. Inner lip thin, simple,
flexed at umbilicall rim, channelled below.

Animal unknown.

Type data. Holotype mnhn (1.35 x

3.45 mm, 4.10 tw) and 62 paratypes (ams, bmnh,
mnhn. nmnz, nmp, usnm) : “ Vauban ", stn 40.
Paratypes (4 mnhn) : Biocal, stn DW 77.

Distribution. OIT Southern New Caledo-

nia, 250-440 m (dead).

Remarks. Fluxinella membranacea differs
from ail hitherto named species of Fluxinella in
the combination of smooth spire, flattened sum¬
mit, persistent shoulder angulation, and dis-
tinctly overhung umbilical wall.

Fluxinella xysila sp. nov.

Figs 98, 101, 103 ; Table 12

Description. Shell up to 6.40 mm, mar-
kedly broader than high, sublenticular, thin,
glossy. umbilicate, spire 0.91-1.17 x as high as
aperture ; white, nacreous through thin, translu¬
cent shell layer.

Protoconch 350-370 (i.m wide, smooth.

Table 12. Fluxinella xysila. Shell measurements (mm) and
countings.

H

D

H/D

TW

UD%

2.10

5.30

0.40

4.30

31.7

Holotype

2.45

6.40

0.38

4.40

31.3

Para type

2.45

6.30

0.39

4.50

30.2

Paratype

Teleoconch of up to 4.50 whorls. First 2 whorls
weakly convex, a weakly defined shoulder angu¬
lation on Ist half whorl, later whorls almost fiat,
peripheral keel prominent, sharp, flange-like]
smooth ; base weakly convex. A crisp spiral
thread bordering umbilicus, entire surface other-
wise smooth apart from fine collabral growth
fines. Umbilicus deep, wall steeply tapered, dia¬
meter 30.2-31.7% of shell diameter. Aperture
subrhomboidal. Outer lip rim damaged, thin

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

69

Figs 91-105. Genus Fluxinella : 91. 93. 94. Fluxinella membranacea, holotypc. 1.35 x 3.45 mm
pni-atypc. " Vauban " stn 40. x 105. 92. 95, 96. 100. 105, F. polita, holotypc. 1,90 x 4.10 m

99, 102. 104, F. brychia, holotypc. 2.10

98. 101, 103, F. xysila , 2.10 x 5.20 mm, 98 x 73.

97. F. membranacea.
). 96 x 30, 100 x 95.
< 5.20 mm. 99 x 70.

Source : MNHN, Paris

70

BRUCE A. MARSHALL

not, significantly thickened within, from growth
Unes posterior notch retraction and protraction
depths 3.7 % of shell diameter ; basal notch
shallow. concave, peripheral notch within keel.
Pariétal glaze thin. Inner lip thin, simple.

Animal unknown.

Type data. Holotype (2.10 x 5.20 mm.
4.20 tw) and paratype (2.45 x 6.40 mm. 4.40 tw)
mnhn, paratype nmnz : Biocal, stn DW 106.

Distribution. — Off Southern New Caledo-
nia, 625-650 m (dead).

Remarks. — Fluxinella xysila appears to be
most closely related to the New Zealand species
F. lenliculosa (803-846 m), from which it differs
in having a higher spire and a more weakly
convex base. Other similar taxa are F. dis¬
cuta (Dali, 1889) (northwestern Atlantic, I 597-
1796 m), F. vitrea (Okutani, 1968) (Japan,
2 100 m), F. gellida (Barnard, 1963) (South
Africa, 2 268-2 377 m) and F. lepida Marshall,
1983 (New Zealand, 1 457-1 463 m). F. xysila
differs from F. gellida in having a much more
weakly convex base, and from the others in
having a more weakly defined shoulder angula¬
tion on the first teleoconch whorl. It differs
further from F. discuta and F. vitrea in having a
smaller protoconch (diameter 350-370 gm cf.
400 u.m), from F. vitrea in having a fiat rather
than weakly convex umbilical wall, and from F.
lepida in having a steeply tapered, rather than
vertical umbilical wall.

Etymology. — Smooth (Greek).

Fluxinella hrychia sp. nov.

Figs 99, 102, 104

Description. — Shell up to 5.20 mm wide,
considerably broader than high, sublenticular,
thin, glossy, umbilicate, spire shallowly cyrtoco-
noid. 0.64-0.95 x as high as aperture ; white,
nacreous through thin, translucent outer shell
layer.

Protoconch 370 ji.m wide, smooth.

Teleoconch of up to 4.2 whorls. First whorl
with supramedian shoulder angulation, sharp at
first, progressively weakening, becoming obsolète
early on 2nd half of lst whorl. First 2 spire

whorls weakly convex, subséquent spire whorls
and base weakly convex, shallowly concave
above and below very prominent, smooth, sharp
peripheral keel. Smooth apart from flexuous
collabral growth fines, and obscure spiral fines
on base, a spiral thread at umbilical rim. Umbili-
cus very broad, deep, rim sharply angulate, wall
steeply tapered, diameter 32.7-35.3 % of adult
shell diameter. Aperture subquadrate. Outer lip
thin, rim damaged, from growth fines posterior
notch concave, retraction depth 2.6 % and pro¬
traction depth 7.7 % of shell diameter ; periph¬
eral notch within keel, not retracted ; basal notch
concave. Pariétal glaze very thin. Inner lip thin,
simple.

Animal unknown.

Type data. - Holotype mnhn (2.10 x
5.20 mm, 4.2 tw) : Biocal, stn CP 72. Paratype
(1 mnhn) : Biocal, stn DS 59.

Distribution. Off Southern New Caledo-
nia, 2 100-2 650 m (dead).

Remarks. Fluxinella brychia bears a strong
general similarity to F. xysila sp. nov. and the
following species with which it is compared. It
differs from F. discuta in having the whorls more
weakly stepped within the umbilicus and in
having a shallowly cyrtoconoid instead of rather
evenly conical spire, from F. discuta and F. vitrea
in having a smaller protoconch (diameter 370 |*m,
cf. 400 jim), and from F. vitrea in having a fiat
rather than weakly convex umbilical wall. It
differs from F. lenliculosa in having a higher
spire, and a shoulder angulation on the first
teleoconch whorl. and from F. xysila in having a
weaker spiral thread at the umbilical rim, a more
sharply defined shoulder angulation, and a deeper
posterior notch. The base is more weakly convex
than in F. gellida.

Etymology. From the deep (Greek).

Fluxinella polita sp. nov.

Figs 92, 95, 96, 100, 105; Table 13

Description. Shell up to 4.40 mm wide,
considerably broader than high, sublenticular
stout, glossy, umbilicate, spire 1.31-1.69 x a g
high as aperture ; white. nacreous through thin
translucent outer shell layer.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

71

Table 13. Fluxinella poliia. Shell measuremenls (mm)
and countings. (Biocal. stn DW 44).

Character

n

Range

Mean

SD

H

10

1.75-2.15

1.86

0.12

D

10

3.65-4.40

3.77

0.98

H/D

10

0.42-0.52

0.46

0.03

TW

10

4.25-4.75

4.51

0.17

UD%

10

18.2-21.7

20.0

1.29

Protoconch 300-330 jxm wide, smooth apart
from faint axial wrinkles at summit.

Teleoconch of up to 4.75 whorls. Spire whorls
fiat or very weakly convex above very promi¬
nent, sharp, thin, flange-like peripheral keel,
which is shallowly concave above, and fiat and
almost horizontal below, drooping abapically at
end of last adult whorl. Whorls smooth apart
from fine collabral growth fines and obscure
spiral fines, umbilical rim rendered shallowly
undulant by weak, bordering radial pleats. Umbili-
cus deep, rim narrowly rounded, projecting
inwards to overhang wall, diameter 18.2-21.7 %
of adult shell diameter. Aperture trapezoidal.
Outer lip thin at rim of labial projection and
base, thick at inner basal extremity. thickened
within, strongly so against posterior notch and
peripheral keel ; posterior notch shallow, apical
rim gently flared, retraction depth 2.38-2.50 %
and protraction depth 5.82-7.55 % of shell dia¬
meter ; basal notch concave ; peripheral notch
within keel, not retracted. Pariétal glaze thin.
Inner lip very thick, concave, flexed at base to
form small, rounded tooth, below which is a
narrow, shallow groove.

Animal unknown (dried).

Type data. — Holotype (1.90 x 4.10 mm,
4.7 tw) mnhn, and 755 paratypes (ams, bmnh,
MNHN, NMNZ, nmp, usnm): Biocal, stn DW 44.

Other material examined (12 specimens mnhn).
— Biocal, stn DW 08 (2). Stn DW 46 (7).

Stn DW 53 (1). Stn DW 70 (2).

Distribution. Off Ouvéa, Loyalty Is, and
Southern New Caledonia, 435-1 005 m, living at
440-610 m.

Remarks. — Fluxinella polila difTers from
hitherto known species of Fluxinella in its stout

glossy shell, smooth periphery, toothed inner lip,
and radially pleated umbilical rim.

Etymology. Made smooth (Latin).

Fluxinella runcinata sp. nov.

Figs 106, 108, 110; Table 14

Description. — Shell up to 4.65 mm wide,
considerably broader than high, stout, glossy,
umbilicate; spire weakly cyrtoconoid, 1.05-1.30 x
as high as aperture ; white, nacreous through
translucent outer shell layer.

Protoconch 330-350 |xm wide (usually 330 (j.m),
smooth.

Table 14. Fluxinella runeinata. Shell measurements (mm)
and countings. (Biocal. stn DW 48. DW 53).

Character

n

Range

Mean

SD

H

10

2.00-2.68

2.33

0.24

D

10

3.75-4.65

4.16

0.31

H/D

10

0.53-0.59

0.55

0.02

TW

10

4.50-5.10

4.84

0.19

UD%

10

21.5-26.1

23.9

1.38

Teleoconch of up to 5.10 whorls. Spire whorls
fiat above shallowly concave adapical side of
prominent, smooth, sharp peripheral keel, base
very weakly and rather evenly convex from
periphery to umbilical rim. Spire whorls smooth
apart from fine collabral growth fines. Base with
2-4 fine, close, similar spiral threads near periphery.
another crisply defincd thread at umbilical rim,
elsewhere smooth apart from collabral growth
fines and obscure spiral fines. Umbilicus deep,
rim smooth. narrowly rounded, weakly projecting
inwards to slightly overhang wall, diameter 21.5-
26.1 % of adult shell diameter. Aperture subtra-
pezoidal. Outer lip thin at rim of labial projec¬
tion and base, moderately thickened within
against posterior notch and peripheral keel ;
posterior notch shallow, broad, retraction depth
2.59-3.54 % and protraction depth 1.30-4.45 %
of shell diameter ; basal notch concave ; periph¬
eral notch within keel, not retracted. Pariétal
glaze thin. Inner lip thick, concave, flexed at base
to form small rounded tooth, below which is a
narrow, shallow groove.

Animal unknown.

Source : MNHN, Paris

72

BRUCE A. MARSHALL

Figs 106-120. — Genus Fluxinella : 106, 108,110, Fluxinella runcinaia, holotype, 2.05 * 3.85 mm. 110 x 90. 107, 109 115

F. asceta, paralypc, Biocal stn DW 33. 3.80 x 6.50 mm. 115 x 80. 111-114, F. asceta, holotype. 4.00 x 6.90 mm’

114 x 20. 116-119, F. megalomphala, holotype, 4.05 x 7.80 mm, 119 (last rw) x 25. 120, F. megalomphala

paratype. Biocai. stn DW 80, x 70.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

73

Type data. Holotype (2.05 x 3.85 mm,
4.6 tw) and 2 paratypes mnhn : Biocal,
stn DW 48. Paratypes (31) : Biocal, stn DW 49
(2 mnhn). Stn DW 53 (1 ams, 1 bmnh,
6 mnhn, 2 nmnz, 1 nmp, 1 usm). — Stn DW 70
(Il mnhn). Stn DW 79 (2 mnhn). —
Stn DW 80 (4 mnhn).

Distribution. OfT Ouvéa, Loyalty Islands,
and southem New Caledonia, 715-1 380 m (dead).

Remarks. Compared with Fluxinella poli ta ,
which it most resembles, F. runcinata differs
primarily in its higher spire and narrower periph-
eral keel, and in lacking radial pleats at the
umbilical rim. The two species hâve overlapping
bathymétrie ranges and they occurred together at
Biocal stations DW 53 and DW 70.

Etymology. Planed off (Latin).

Fluxinella asceta sp. nov.

Figs 107, 109, 111-115, 280, 281; Table 15

Description. Shell up to 7.20 mm wide,
markedly broader than high, trochiform, stout,
glossy, umbilicate, spire shallowly cyrtoconoid,
1.74-2.29 x as high as aperture ; white, nacreous
through thin, translucent ouler shell layer.

Table 15. Fluxinella asceta. Shell measurements (mm)
and countings. (Biocal, stn DW 33).

Character

n

Range

Mcan

SD

H

10

3.00-4.55

3.77

0.49

D

10

5.33-7.20

6.49

0.65

H/D

10

0.53-0.63

0.58

0.03

TW

10

6.00-7.00

6.47

0.26

UD%

10

20.8-24.4

22.5

1.18

Protoconch 270-300 ^m wide, smooth, tip excert.

Teleoconch of up to 7 whorls. Shoulder angula¬
tion strong on lst whorl, progressively weakening,
vanishing on next whorl. Subséquent whorls
shallowly concave, adapical two-thirds fiat, aba-
pical third weakly convex at first, becoming fiat
and less steeply sloping than adapical two-thirds
or grading to shallowly concave. Peripheral keel
sharp-edged, narrowly angulate in section, strongly
projecting, summit very weakly undulant, basal
side more or less horizontal, keel often distinctly

drooping at maturity, occasionally slightly up-
turned. Base weakly convex. Axial riblets on
spire low, rounded, collabral, shallowly sigmoidal.
resolving from weak undulations that commence
on 2nd whorl, most strongly defined over abapical
third, becoming strongly defined over adapical
two-thirds on last part of last adult whorl. Spire
whorls at first with single spiral thread at summit
of shoulder angulation, becoming obsolète on
about mid 3rd whorl, 2nd spiral commencing at
end of 2nd whorl between shoulder spiral and
periphery, tending to persist throughout at aba¬
pical third. a 3rd spiral often appears after 4th
whorl between 2nd spiral and periphery. Spire
whorls otherwise smooth apart from fine colla¬
bral growth Unes and obscure spiral Unes over
adapical two-thirds. Base with 2-4 crisp spiral
threads on outer part, and strong rounded axial
pleats at umbilical rim, otherwise smooth apart
from collabral growth fines and obscure spiral
fines. Umbilicus deep, wall shallowly concave,
steeply tapered, diameter 20.8-24.4 % of adult
shell diameter. Aperture subtrapezoidal. Outer
lip rather thin at rim, modestly and rather
uniformly thickened within ; posterior notch
broad, concave, retraction and protraction
depths, 2.4 % of shell diameter ; basal notch
broad, concave ; peripheral notch within keel,
not retracted. Pariétal glaze thin. Inner lip thick,
deeply curved towards umbilicus, gently flexed at
base.

Animal. Snout at least twice as long as broad,
latéral projections at tip rounded, mouth a
vertical slit beneath. Cephalic tentacles about
twice as long as snout, dorsoventrally flattened,
narrow, very gradually tapered, inner bases at
least 2 tentacle base-widths apart. edges ciliate,
large swollen black eyes at outer bases. Right
suboptic tentacle large, shorter and stouter than
cephalic tentacles, gradually tapered, tip rather
blunt, ventrally grooved. Epipodial tentacles
numbering 9 on right and 1 or 2 on left, right
tentacles decreasing in size towards opercular
lobe, left tentacles small and short. Operculum
chitinous, spiral.

Radula (Figs 280, 281) with the formula
c. 20 + 1 + 1 + 1 + c. 20. Central tooth rigid,
slightly longer than broad, cutting area jutting
forward at right angle from shaft, broadly
angulate, with about 9 stout cusps, médian cusp
largest, latérobasal projections prominent. Laté¬
ral teeth broad, cutting area of each angulate,

Source : MNHN, Paris

74

BRUCE A. MARSHALL

terminal cusp largest, finer cusps on outer edge,
fewer on inner edge. Innermost marginal broader
than outer marginals, cutting area narrowly
angulate, terminal cusp large, finer cusps on
outer edge, a few fine cusps on inner edge. Outer
marginals slender, each wilh small, slender cusps
at tip and on outer edge.

Jaw plates subrectangular, thin, broader than
long, éléments short, longer anteriorly.

Type data. Holotype mnhn (4.00 x
6.90 mm, 6.70 tw) and 193 paratypes (ams,

BMNH, MNHN, NMNZ, NMP, USNM) : BlOCAL,

stn DW 33.

Other material examined (125 specimens mnhn).

Biocal, stn DW 36 (4). — Stn DW 46 (8).

Stn DW 48 (19). — Stn DW 51 (81). — Stn DW 53
(7). — Stn DW 70 (2). — Stn DW 80 (4).

Distribution. OIT Ouvéa, Loyalty Islands,
and Southern New Caledonia, 570-1 005 m, living
at 570-700 m.

Remarks. Fluxinella asceta is highly dis¬
tinctive in its large size, tall spire, low widely
spaced axial riblets, and crisp peripheral spiral
threads.

Etymology. Ornamented (Greek).

Fluxinella megalomphula sp. nov.

Figs 116-120

Description. — Shell up to 7.80 mm wide,
considerably broader than high, depressed tro-
chiform, stout, rather thin, spire 1.41-1.43 x as
high as aperture ; white, nacreous through thin,
translucent outer shell layer.

Protoconch 370 ;xm wide, essentially smooth.

Teleoconeh of up to 6.30 whorls. Shoulder
angulation sharp on lst whorl, weakening and
vanishing on 2nd whorl ; subséquent whorls
grading from fiat to weakly convex above promi¬
nent sharp, flange-like peripheral keel. Base
suddenly contracted, weakly convex. Axial riblets
commencing on 3rd whorl, widely spaced, low,
rounded, weakly flexuous, rendering periphery
weakly undulant. Base with 5 spiral threads on
outer third, 3 or 4 spiral cords on inner quarter,
and rounded, fold-like axial costae that résolve

on inner half and enlarge towards umbilicus,
extending onto outer part of umbilical wall.
Additional spiral cords resolving on broad médian
basal zone on last half adull whorl. Umbilicus
deep, wall steeply tapered, diameter 32.3-35.9 %
of adult shell diameter. Aperture subrhomboidal.
Outer lip thin at rim, thickcr within, posterior
notch shallow and broad, retraction depth (from
growth lines) 1.70-2.00 %, and protraction depth
2.10-2.60% of shell diameter; basal notch con¬
cave, peripheral notch within keel. Pariétal glaze
thin. Inner lip thick, curved towards umbilicus
adapically, flexed below to form small, broadly
rounded submedian projection.

Animal unknown.

Type data. Holotype (4.05 x 7.80 mrn ,
6.30 tw) and paratype mnhn ; paratype nmnz
(3.40 x 6.50 mm, 6.00 tw) : Biocal, stn DW 80.

Distribution. OIT Ouvèa, Loyalty Islands,
900-980 m (dead).

Remarks. — Compared with Fluxinella asceta,
to which it is superficially similar, F. megahm-
phala differs principally in having a more broadly
conical spire, finer spiral threads at the periph¬
ery, and a much wider umbilicus. The two
species occurred together at the type locality.

Etymology. Big navel (Greek).

Fluxinella euphanes sp. nov.

Figs 121-125

Description. — Shell up to 4.65 mm wide,
considerably broader than high, sublenticular,
thin, spire 0.94-1.05 x as high as aperture;
white, nacreous through thin, translucent outer
shell layer.

Protoconch 350-370 gm wide, minutely granu-
late.

Teleoconeh of up to 4.90 whorls, periph¬
ery sharply angulate, flange-like ; base sud¬
denly contracted, outer third below keel more or
less horizontal, fiat or shallowly concave, inner
part convex. Shoulder angulation strong on
lst whorl, progressive^ weakening, vanishing
early on 3rd whorl, commencing level with
suture, descending to about adapical third, as-
cending late on 2nd whorl ; ramp and side

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZ1IDAE

75

Figs 121-134. Gcnus Fluxinella : 121-125, Fluxinella euphanes, hololype, 1.90 * 4.65 mm, 124 (laie 3rd tw) x 60,
125 x 70. 126-129, F. tenera , holotype, 2.30 x 4.60 mm, 129 (last TW) x 60. — 130-134, F. stirophora, hololype.

2.10 x 4.10 mm. 133 (last tw) x 50. 134 x 95.

Source : MNHN, Paris

76

BRUCE A. MARSHALL

concave. Third and later whorls with flattened
sub and suprasutural zones, subsutural zone
depressed, progressively grading from about third
to half whorl height, becoming steeper than
raised suprasutural zone. Spire whorls with
prominent, widely spaced, shallowly flexuous
axial riblets that render periphery gently undu-
lant. Base with 3 or 4 fine, crisp spiral threads on
outer third, and prominent, rounded axial pleats
bordering umbilicus ; elsewhere smooth apart
from collabral growth fines and obscure spiral
fines. Umbilicus deep, wall steep, weakly convex,
diameter 25.0-25.4 % of adult shell diameter.
Aperture subrhomboidal. Outer lip rim damaged,
from growth fines posterior notch shallow and
broad, retraction depth 2.20 % and protraction
depth 5.7 % of shell diameter : basal notch
concave, peripheral notch within keel. Pariétal
glaze thin. Inner lip hollow and comprising 2
thin parallel walls adapicaliy, flexed submedially
to form small, rounded tubular tooth ; rim
becoming covered over at maturity, thin and
simple below tooth.

Animal unknown.

Type data. Holotype (1.90 x 4.65 mm,
4.90 tw) and paratype (1.60 x 4.13 mm, 4.80 tw)
mnhn : Biocal, stn DW 79.

Distribution. Off Ouvéa, Loyalty Islands,

I 320-1 380 m (dead).

Remarks. Among previously described
species of Fluxinella, F. euphanes is very distinc¬
tive in the stepped contour of the last adult and
the strong, widely spaced axial riblets. The
double or at least locally hollow inner lip wall in
this and the following taxon is a most unusual
character hitherto unknown from the family (see
also Quinnia limatula sp. nov.).

Etymology. Very bright (Greek).

Fluxinella tenera sp. nov.

Figs 126-129

Description. — Shell (holotype) 4.60 mm
wide, considerably broader than high, thin, spire
1.08 x as high as aperture ; white, nacreous
within.

Protoeonch eroded, about 300 [un wide.

Teleoconch of 4.8 whorls, periphery sharply

angulate, flange-like ; base very suddenly con-
tracted below periphery. convex. First 1.5 whorl
eroded, next whorl shallowly concave below low
subsutural bulge, subséquent whorls divided into

2 flattened zones, zone at abapical half slightly
raised, shallowly concave on last adult whorl.
Spire whorls entirely traversed by narrow, widely
spaced, sigmoidal, collabral axial riblets. Aba¬
pical médian spiral commencing on 2nd half of
3rd whorl, adapical médian spiral commencing a
whorl later, gradually enlarging to resemble
abapical spiral ; a weak secondary spiral com¬
mences at end of 4th whorl between abapical
médian spiral and periphery, vanishing within
half a whorl. Spire whorls rendered dull by
minute granules and obscure spiral fines. Outer
base with 4 crisp spiral threads on peripheral
flange, innermost 2 weaker. Inner base with

3 rounded spiral cords, innermost at umbilical
rim ; and rounded, fold-like axial riblets that
résolve midway across base and enlarge towards
umbilicus, extending around umbilical rim. Base
with addition of minute granules, obscure spiral
fines, and weak collabral growth fines, the latter
most conspicuous on peripheral keel. Umbilicus
deep, rim rounded to overhang minutely granu-
late, subvertical wall, diameter 22.8 % of shell
diameter. Aperture subtrapezoidal, rim damaged.
Outer lip simple within, posterior notch concave,
from growth fines retraction depth 2.90 % and
protraction depth 6.50 % of shell diameter ;
peripheral notch within keel, basal notch con¬
cave. Pariétal lip thin. Inner lip thick, rim
comprising inner and outer walls that are covered
over at maturity, shallowly concave, produced
and retracted at base to form low, rounded keel
that borders narrow, concave basal channel.

Animal unknown.

Type data. Holotype mnhn (2.30 x
4.60 mm, 4.8 tw) : Biocal, stn DS 14.

Distribution. East of Ouvéa, Loyalty
Islands, 3 680-3 700 m (dead).

Remarks. Compared with Fluxinella eupha¬
nes, which il most resembles, F. tenera diflers
primarily in having two médian spiral cords on
adult whorls, stronger basal spiral sculpture, an
inwardly projecting umbilical rim, and a shallowly
concave rather than convex umbilical wall.

Etymology. Délicate (Latin).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

77

Fluxinella stirophora sp. nov.

Figs 130-134

Description. Shell up to 4.20 mm wide,
rather thin and fragile, umbilicate ; spire low,
conical, 0.77-1.14 x as high as aperture ; white,
nacreous through thin, translucent outer shell
layer.

Protoconch 300 pm wide, surface obscurely
and finely granulate.

Teleoconch of up to 5.25 whorls. First spire
whorl weakly convex at lirst, grading to shal-
lowly concave, 2nd and 3rd whorls shallowly
concave, subsequently grading from concave
to convex though remaining shallowly concave
between suprasutural spiral and periphery ; pe-
riphery angulate, rendered shallowly serrate by
axial riblets, base convex. Axial sculpture on
spire consisting of fine, low. widely spaced,
sigmoidal riblets. A suprasutural spiral thread
commencing immediately, strong al first, progres-
sively weakening, vanishing at end of lst whorl.
A second fine suprasutural spiral thread at
about abapical third commencing at about mid
4th whorl, crisp throughout ; other fine threads
commencing after 4th whorl and multiplying by
intercalation, most crisply defined on abapical
half of last adult whorl. Minute, crowded gra¬
nules throughout. Base with a fine outer spiral
thread, a very broad médian zone that is smooth
apart from fine collabral growth fines, and 3 or
4 inner spiral cords, the innermost strongest,
smooth and separated by a conspicuous groove.
Basal axial riblets confined to zone between
periphery and outermost spiral thread, and
between broad médian zone and inner groove.

Broad médian zone and summits of inner basal
spirals glossy, elsewhere roughened by minute
granules. Umbilicus deep, rim angulate, diameter
23.6-23.8 % of adult shell diameter. Aperture
ovate. Outer lip thin, not thickened within.
mature rim damaged ; from growth fines pos-
terior notch concave, retraction depth 4.9 % and
protraction depth 8.1 % of shell diameter ; basal
notch concave, no peripheral notch. Pariétal
glaze thin. Inner lip short, very thick, a strong.
blunt, rounded denticle at base.

Animal unknown.

Type data. — Holotype mnhn (2.10 x
4.10 mm. 5.25 tw) and paratype nmnz (2.00 x
4.20 mm. 4.80 tw) : Biocal. stn DW 56. Para-
type (mnhn) : Biocal, stn DW 51.

Distribution. — Off Southern New Caledo-
nia, 694-705 m (dead).

Remarks. — Fluxinella stirophora is highly
distinctive among Fluxinella species in the com¬
bination of low spire, convex adult spire whorls,
fine reticulate teleoconch sculpture, grooved umbi-
lical rim, and deeply retracted posterior notch. F.
stirophora is referred to Fluxinella because of its
general resemblance to F. euphanes sp. nov. and
F. tenera sp. nov. This placement is provisional,
however, for when animais are available for
comparison it may prove to belong in tribe
Seguenziini, perhaps Quinnia, species of which
hâve similarly shaped posterior notches and
somewhat similar sculpture.

Etymology. Keeled (Greek).

Genus BASIL1SSA Watson, 1879

Basilissa Watson. 1879 : 593. Type species (by sub¬
séquent désignation of Cossmann, 1888) : Basilissa
superha Watson. 1879; Recent, Coral Sea, east of
Cape York.

Remarks. Although a number of species
hâve been referred to Basilissa since its introduc¬
tion, none are currently regarded as being conge-
neric with B. superha (Quinn, 1983b, 1987). It
seems highly likely, however, that the Atlantic

type species of Thelyssa Bayer, 1971 (T. callisto
Bayer, 1971) belongs here.

Basilissa superha Watson, 1879
Figs 135, 142, 143, 284-287

Basilissa superha Watson, 1879 : 598 ; 1886 : 101. pl. 7.
fig. 10. Cernohorsky, 1978 : 33, pl. 8, fig. 4.

Source : MNHN, Paris

78

BRUCE A. MARSHALL

Figs 135-148. Généra Basilissa, Halyslina. Hadroconus. Carenzia : 135 , 142 , 143 , Basilissa superba (juvénile) Biocai stn DS
59, 4.30 x 4.35 mm. 142 (last tw) x 30. 143 x 45. 136 , 138 , Halyslina siherutensis, lectolypc, 2.40 x 2,(8) mm 138

(laie 3rd tw) x 55. 137 , //. siherutensis, paralectotypc, widlh 1.90 mm. 139 - 141 , Hadroconus grandiosus

hololypc. 9.00 x 11.0 mm. 141 (last tw) x 13. 144 - 148 , Carenzia nitens, holotypc, 2.50 x 2.60 mm, 147 (last tw) x

50. 148 x 70. (For various reasons Figs. 136-138 could not bc includcd in systcmatic séquence).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

79

Description (supplementary to the original).
Spire outline at first shallowly cyrtoconoid,
becoming shallowly coeloconoid.

Proloconch globular, perfectly smooth, 530-
600 [xm wide, apertural rim flared.

Teleoconch : lst whorl essentially smooth
apart from peripheral keel, which is strong
throughout. Axial riblets and traces of spiral
threads gradually resolving late on lst whorl,
axials becoming clearly defined on 2nd whorl,
spirals poorly defined on lst 3 whorls, that
beside peripheral keel enlarging more rapidly,
becoming as strong as axials on 4th whorl, after
which other spirals become as large.

Animal unknown (decayed).

Radula (Figs 284-287) whith the formula
c. 12 + 1 + 1 + 1 +c. 12. Central tooth subrec-
tangular, longer than broad, rigid, cutting area
curving forward, angulate, médian cusp strong,
accessory cusps fine, 6-8 on each side, latérobasal
projections weak. Latéral teeth broad, rigid,
cutting area of each angulate, terminal cusp
strong, accessory cups fine, about 18 on outer
edge and 1 or 2 on inner edge. Innermost
marginal broadest, thin in section, cutting area
narrowly angulate, terminal cusp large, long
sériés of fine accessory cusps on outer edge,
fewer on inner edge. Outer marginals slender,
cusps fine, narrowly conical, long sériés on outer
edge, few on inner edge. Basal plate of each
marginal flanged and grooved to interlock with
laterals and with each other.

Jaw plates thin, subrectangular, considerably
broader than long, éléments short, longer ante-
riorly.

Type data. Holotype bmnh 1887.2.9.354,
Coral Sea, east of Cape York, Queensland,

2 560 m.

Material oxamined (5 specimens mnhn). Bio¬
cal. stn CP 13 (1 adult). — Stn CP 17 (1 adult,
1 subadult). — Stn DS 59 (2 juvéniles).

Distribution. Northern Coral Sea (2 560 m),
off Lifou, Loyalty Islands (3 680-3 740 m) and
ofT Southern New Caledonia (2 560 m), living at

3 690-3 740 m.

Remarks. The présent specimens agréé well
with the holotype. Judging from the description
and illustrations (Okutani, 1982), specimens
from the Philippine Sea (3 210-3 680 m) are also
similar, but difFer in having the umbilicus partly
closed by a septum. The Coral Sea and Philip¬
pine Sea populations are separated by island arcs
and deep trenches, and the status of the Philip¬
pine Sea form is uncertain. The jaw plates, which
disintegrated during cleaning, are considerably
broader relative to length than in Ancistrobasis,
Calliobasis and Fluxinella. Convoluted inter-
locking marginal basal plates hâve not been
hitherto recorded from the family. but since the
bases of the marginal teeth of other seguenziids
hâve not been studied, their significance is
uncertain. This aptly named species is the largest
seguenziid known, the largest of the présent
specimens (Biocai. stn CP 17) having a diameter
of 17.7 mm and an estimated height of 21 mm.
See Discussion page 107.

Tribe SEGUENZ1INI
Genus HADROCONUS Quinn, 1987

Hadroconus Quinn, 1987 : 61. Type species (by
original désignation) : Basilissa alla Watson, 1879 ;
Recent, Atlantic central America.

Hadroconus grandiosus sp. nov.

Figs 139-141

Description. Shell (holotype) 11.0 mm
wide, broader than high, thin, umbilicate, spire

1.75 x as high as aperture ; white, nacreous
through thin, translucent outer shell layer.

Proloconch 400 um wide, surface eroded.

Teleoconch of 7.8 whorls; lst whorl convex,
subséquent whorls flattened. suture flush. periph-
ery angulate, base suddenly contracted, weakly
convex. Axial riblets fine, crisp. opisthocline and
noncollabral on spire ; sigmoidal, collabral and
much weaker on base, though forming promi¬
nent, rounded, radial pleats at umbilical rim.

Source : MNHN, Paris

80

BRUCE A. MARSHALL

Spire spirals multiplying by intercalation, absent
from broad médian zone on 3rd-6th whorl, after
which evenly distributed over adapical three
quarters. Spiral at abapical quarter strongest,
angulate in section, with small conical nodules at
intersections with axials ; other spirals similar in
size, thread-like. Basal spiral cords increasing in
number to 18; outer 5 similar, angulate, inter-
spaces broader than each spiral ; inner spirals
flattened, interspaces narrower than each spiral.
Umbilicus deep, diameter 25.4 % of shell diamc-
ter. Aperture trapezoidal. Outer lip thin, slightly
thickened within ; posterior notch broad, rim
damaged, retraction depth 3.6 % of shell diame¬
ter, protraction depth unknown (though cer-
tainly at least 11 % of shell diameter) ; basal
notch shallow, concave ; peripheral notch within
peripheral angulation, not retracted. Pariétal
glaze thin. Inner lip rather thick, rim tightly
folded towards umbilicus, curved towards umbi¬
licus at insertion, almost straight below, tooth-
less.

Animal unknown.

Type data. Holotype mnhn (9.00 x

11.00 mm, 7.8 tw) r Biocal, stn CP 57.

Distribution. South of New Caledonia

I 490-1 620 m (dead).

Remarks. Compared with the closely similar
H. al tus (Watson. 1879) (north-western Atlantic,
c. 500-2 360 m), //. grandiosus diflers primarily in
having considerably doser, more numerous axial
riblets thaï form weaker nodules at the peri-
phery. It diflers from //. sibogae (Schepman,
1908) (Indonesia, 1 158-1 301 m) in having more
numerous spiral threads on the spire and weaker
axials on the base, and from //. diadematus
Marshall, 1988 (New Zealand, I 463-1 457m) in
having fewer spiral threads on the spire before
the last adult whorl, and in being smaller relative
to the number of whorls.

Etymology. Enlarged (Latin).

Genus CARENZ/A Quinn, 1983

Carenziu Quinn, 1983 : 355. Type species (by original
désignation) : Seguenzia carinata JefTreys, 1877 ;
Recent, North Atlantic.

Carenzia nitens sp. nov.

Figs 144-148

Description. Shell up to 2.62 mm wide,
about as broad as high, thin, umbilicate, spire
1.15-1.28 x as high as aperture ; white, nacreous
through thin, translucent outer shell layer.

Protoconch 270-300 p.m wide, with 6 fine spiral
threads that vanish before apertural rim.

Teleoconch of up to 4.9 whorls. shoulder and
peripheral angulations smooth ; ramp liât at
first. becoming weakly convex ; side shallowly
concave throughout or becoming fiat ; base
suddenly contracted, convex. Shoulder angula¬
tion supramedian, sharp, not projecting, becom¬
ing obsolète late on last adult whorl. Peripheral
keel strong, sharply angulate. Base with weak
spiral cord near periphery, obscure spiral fines,
and rounded spiral cord at umbilical rim. First
whorl finely and sparsely granulate. Fine colla-

bral axial riblets commence late on lst whorl,
crisply defined on ramp, weaker on side, weak-
ening and vanishing on lst half of 3rd whorl.
Subséquent whorls olherwise smooth apart from
fine collabral growth fines. Umbilicus deep,
diameter 16.7-28.6 % of adult shell diameter.
Aperture subrhomboidal. Outer lip thin, simple
within ; posterior notch retraction depth 5.11 %
and protraction depth at least 15% of shell
diameter (tip of labial projection broken) ; basal
notch shallower, peripheral notch weak. Pariétal
glaze thin. Inner lip curved toward umbilicus,
flexed near base to form small rounded tooth.

Animal unknown.

Type data. Holotype mnhn (2.50 x
2.60 mm, 4.5 rw) and 2 paratypes mnhn, nmnz
( 2.40 x 2.40 mm, 4.6 tw ; 2.20 x 2.62 mm,
4.9 tw) : Biocal, stn DW 79. Paratypes
(3 mnhn) : Biocal, stn CP 57.

Distribution. OU Ouvéa, Loyalty Islands,
and Southern New Caledonia, 1320-1 620 m
(dead).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

81

F,gs >49- 63. Genus Carenzia : 149-153, Carenzia serrata, holotype. 2.85 x 3.45 i
154-157, C. acanthodes. hololypc. 3.95 x 5.10 mm, 157 (last tw) x 30. 158,

26, x 70. 159-163, C. urnata, holotype, 2.70 x 3.60 mm, 162 (last TW)

î, 152 (last tw) x 40. 153 x 100. —
. acanthodes, paratype. Biocal stn CP
40. 163 x 80.

Source : MNHN, Paris

82

BRUCE A- MARSHALL

Remarks. Carenzia nitens resembles lhe
New Zealand species C. fastigiata Marshall.
1983, and difiers in attaining maturity at smaller
size, and in having a weaker. unpleated peripheral
keel.

Etymology. Shining (Latin).

Carenzia serrata sp. nov.

Figs 149-153

Description. Shell up to 3.45 mm wide,
broader than high, rather thin, umbilicate, spire
1.17 x as high as aperture, glossy ; white, nacreous
through thin, translucent outer shell layer.

Protoconch 270 (j.m wide, very finely granulate
at extreme tip of apical fold, elsewhere smooth.

Teleoconch of up to 5.75 whorls, shoulder and
periphery angulated by sharp keels, ramp shal-
lowly concave at first, becoming weakly convex ;
side concave ; base suddenly contracted, weakly
convex. Shoulder keel strongly supramedian at
first, descending until almost médian, peripheral
keel more strongly projecting ; both smooth at
first, with small, sharp, conical nodules after
3rd whorl. Nodules coinciding with weak axial
riblets between shoulder keel and outermost
basal spiral, axials strongest on shoulder and
peripheral keels, and between peripheral keel and
outermost basal spiral, almost obsolète between
keels and beside outermost spiral. Spire whorls
otherwise smooth apart from fine, sigmoidal,
collabral growth Unes. Base with 11 spiral cords,
the 2 beside outermost spiral ill-defined, inner
9 spirals reticulating with fine collabral axial
riblets. Umbilicus deep, diameler 26 % of adult
shell diameter. Aperture subrhomboidal. Outer
lip thin at rim, modestly thickened within,
posterior notch concave, rim flared, rétraction
depth 8.7% and protraction depth 11.6% of
adult shell diameter ; basal notch narrower, rim
flared ; periphery not retractively notched though
adult rim concave and flared. Pariétal glaze very
thin. Inner lip curved toward umbilicus, flexed
near base to form strongly projecting rounded
tooth.

Animal unknown.

Type data. — Holotype mnhn (2.85 x
3.45 mm, 5.75 tw) : Biocal, stn CP 26. Paratype
(1 mnhn) : Biocal, stn CP 57.

Distribution. Off Southern New Caledo-
nia, 1 490-1 740 m (dead).

Remarks. Compared with C. trispinosa
(Watson, 1879) (Quinn, 1983a, figs 8-12), which
it most closely resembles, C. serrata difiers in
having the shoulder angulation set considerably
lower on the whorls, and in having a smaller
protoconch (diameter 270 [j.m, cf. 380 |xm). It
difiers from the New Zealand species C. fasti¬
giata Marshall, 1983 in having a lower peripheral
keel and prominenl nodules on the shoulder
angulation.

Etymology. Saw-toothed (Latin).

Carenzia acanthodes sp. nov.

Figs 154-158

Description. Shell up to 5.10 mm wide,
markedly broader than high, of moderate thick-
ness. stout, widely umbilicate, spire 1.76 x as
high as aperture ; white, nacreous through thin,
translucent outer shell layer.

Protoconch 330-370 |im wide, tip excerl, spar-
sely and very finely granulate.

Teleoconch of up to 6.10 whorls, shoulder and
periphery angulated by projecting, angulate keels,
ramp more or less fiat, side concave, base
suddenly contracted. weakly convex. Spire with
3 spiral keels. shoulder keel strongly suprame¬
dian at first, descending until almost médian,
keels smooth at first, shoulder and peripheral
keels with conical nodules on 3rd and subséquent
whorls ; subsutural keel gradually resolving from
low, rounded swelling. with conical nodules on
4th and subséquent whorls. Nodules coinciding
with weak collabral axial riblets that gradually
résolve on 2nd whorl, most crisply defined
between shoulder keel and outermost basal spi¬
ral, almost obsolète on ramp. Spire otherwise
smooth apart from obscure spiral fines and fine
sigmoidal, collabral growth fines. Adult base
with 8 subequal spiral cords and weaker colla¬
bral axial riblets. Umbilicus deep, perspective to
protoconch, diameter 32.5-34.0 % of adult shell
diameler. Aperture subrhomboidal. Outer lip
simple within. intact rim unknown, from growth
fines rétraction depth 5.88 % and protraction
depth 6.52 % of shell diameter ; basal notch
shallower, very slightly notched at periphery.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

83

Pariétal glaze very thin. Inner lip rather thick,
curved toward umbilicus. flexed near base to
form small rounded tooth.

AnimaI unknown.

Type data. - Holotype mniin (3.95 x
5.10 mm, 6.10 tw) : Bioc al, stn CP 26. Para-
types (4 mnhn, 1 nmnz) : Biocal, stn DW 79.

Distribution. OIT Ouvéa, Loyalty Islands
and Southern New Caledonia. 1320-1 740 m
(dead).

Remarks. — Carenzia acanthodes closely resem-
bles C. melvilli (Schepman, 1909) (Makassar Strait,
Indonesia, I 301 m) in general faciès and the two
species are undoubtedly closely related congénère.
By direct comparison with the holotype (zma), C.
acanthodes diffère in having a lower, more broadly
conical spire, and a substantially broader umbilicus
(diameter 32.2-34.0 % cf. 27.2 % of shell diame-
ter). Although the two forms are interpreted as
distinct species, il must be admitted that there is
currently insufficient material to assess the limits of
variation in shell morphology. Compared with C.
serrata sp. nov., with which it occurred at Biocal
station CP 26, C. acanthodes diffère in having a
larger protoconch, in attaining larger size, and in
having a more strongly nodular subsutural spiral.

Etymology. — Prickly (Greek).

Carenzia ornata sp. nov.

Figs 159-163

Description. Shell up to 3.60 mm wide
(immature?) markedly broader than high, rather
thin, widely umbilicate, spire up to 1.40 x as high
as aperture ; white. nacreous through thin, translu-
cent outer shell layer.

Protoconch 370-380 ^m wide, surface almost
entirely etched away but intact surface beside
suture apparently smooth.

Teleoconch of up to 5.1 whorls, shoulder and
periphery angulated by sharply angulate keels,
ramp and side concave ; base suddenly contracted,
weakly convex. Shoulder keel strongly supra-

median al first, descending until almost médian,
peripheral keel stronger and more narrowly
angulate. Shoulder and peripheral keels smooth
at first, small conical nodules on peripheral
keel after late lst whorl and on shoulder keel
after 2nd whorl. An angulate subsutural keel
with small conical nodules commences late on
3rd whorl. Summits of keels on last 2 adult
whorls with fine spiral threads, 2 each on sub¬
sutural and peripheral keels, 3 on shoulder keel,
additional threads between periphery and outer-
most basal spiral. Axial riblets fine, crisp, sigmoi-
dal, collabral, commencing late on lst whorl.
Axials at first between shoulder and periphery,
extending adapically onto ramp on lst half of
3rd whorl and progressively extending to suture.
From late 3rd whorl axials continuous over spire
whorls and base, including outer part of umbili-
cal wall. Base either with 9 spiral cords. the outer
3 weaker, or 8 similar spiral cords. Umbilicus
deep, rim sharply angulate, diameter 27.8 % of
shell diameter. Aperture subrhomboidal, intact
rim unknown. From growth Unes retraction
depth 2.77 %, and protraction depth 10.19 % of
shell diameter. Pariétal glaze very thin. Inner lip
(immature ?) thin, shallowly sigmoidal, toothless.

Animal unknown.

Type data. Holotype mnhn (2.70 x
3.60 mm, 5.1 tw) : Biocal, stn CP 72. Paratypes
(2 mnhn) : Biocal, stn DS 59. Stn DS 98.

Distribution. - Off Southern New Caledo¬
nia, 2 100-2 650 m (dead).

Remarks. Carenzia ornata most closely
resembles C. melvilli (Schepman. 1909) and C.
acanthodes sp. nov., differing from the former in
having finer sculpture on the spire and a broader
umbilicus, and from the latter in having a larger
protoconch, a thinner shell, and doser and
stronger axial sculpture. That the présent spéci¬
mens may be immature is suggested by the
exceptionally large protoconch and the simple
inner lip.

Etymology. — Ornate (Latin).

Source ! MNHN, Paris

84

BRUCE A. MARSHALL

Genus QUINNIA Marshall, 1988

Seguenziella Marshall, 1983 : 245. Type species (by
original désignation) : Seguenziella patula Marshall.
1983 ; Recent, New Zealand (not Seguenziella
Neviani, 1901, not Sacco, 1904).

Quinnia Marshall, 1988 : 242. Replacement namc for
Seguenziella Marshall, 1983 (preoccupied).

Quinnia patula (Marshall, 1983)

Figs 164-168; Table 16

Seguenziella patula Marshall, 1983 : 245, figs 4a-e. 8 j-l.
Quinnia patula - Marshall, 1988 : 242.

Type data. Holotype. nzoi h. 377 : P 939,
41"20' S, 166°54.8' E, E slope of Tasman Basin,
off Westport, New Zealand, I 760-1 799 m.
22 April 1980, R. V. “ Tangaroa

Other material examined (3 speeimens mnhn).
Biocal, stn CP 72 (1). Stn DS 98 (2).

Distribution. Off Southern New Caledonia
(dead, 2 100-2 470 m) and off Westport, New
Zealand (alive, 1760-1 799 m).

Remarks. I am unable to detect any
taxonomically significant différences between New
Caledonian and New Zealand speeimens. The
species is extremely similar lo Q. cazioti (Daut-
zenberg, 1925), based on a specimen taken at
2 286 m off Madeira.

Table 16. Quinnia patula. Shell mçasuremenls (mm)
and countings.

H

D

H/D

TW

UD%

5.65

8.40

0.67

6.10

23.4

Holotype

4.60

7.30

0.63

5.75

31.5

Biocal. su

i CP

72

3.90

6.10 (est.)

0.64

5.40

30.3

Biocal, sti

t DS

98

2.00

3.40

0.59

4.00

23.5

Paratype

1.30

2.45

0.53

3.20

28.5

Bioc al, su

i DS

98

Quinnia laetijica sp. nov.

Figs 169-173

Description. Shell up lo 4.20 mm wide,
broader than high, thin, umbilicate, spire about
as high as aperture ; white, nacreous through
thin. translucent outer shell layer.

Protoconch 370 p.m wide, minutely granulate.

Teleoconch of up to 4.70 weakly convex,
shouldered whorls. Shoulder angulation sharp,
supramedian at first, descending to submedian
position, ramp and side shallowly concave. Peri-
pheral angulation similar to shoulder angulation,
base rather gently contracted, convex. Spire
whorls with similar, fine, crisp, sigmoidal, colla-
bral axial riblets, and spiral threads, very minu¬
tely granulate throughout. Axial riblets evenly
developed over spire lo outermost basal spiral,
weaker on base, evanescent on umbilical wall.
Spiral threads mulliplying by intercalation, at
start of last adult whorl numbering 3 or 7 on
ramp and 3 or 4 on side, subsutural spiral
surmounting low angulation that commences on
2nd half of 3rd whorl. Base with 2 or 3 fine outer
spiral threads below periphery, and 9 stronger
cords that enlarge towards umbilicus ; inter-
spaces considerably wider than each spiral, locally
with addition of I or 2 intercalating spiral
threads. Umbilicus deep. rim sharp, diameter
25.6-28.5 % of adult diameter. Aperture sub-
rhomboidal. Outer lip rim damaged, not thick-
ened within ; from growth fines posterior notch
well retracted from suture, apex roundly angu-
late, rétraction depth 4.28 % and protraction
deplh 9.41% of shell diameter; basal notch
rounded ; no peripheral notch. Inner lip thin,
simple.

Animal unknown (dried).

Type data. Holotype (3.02 x 3.90 mm,

4.40 rw) and paratype (3.58 x 4.20 mm. 4.70 tw)
mnhn : Biocal, stn CP 23 (alive).

Distribution. OIT Southern New Caledo¬
nia, 2 040 m (alive).

Remarks. Quinnia laetifiea differs from Q.
patula and Q. cazioti in having a submedian
instead of supramedian shoulder angulation, a
taller spire, stronger spiral sculpture on the base,
and a narrower umbilicus.

Etymology. Gladdening (Latin).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

85

Figs 164-178. Gcnus Quinnia : 164-167, Quinnia panda. Biocal stn CP 72,4.60 x 7.30 mm, 167 (last tw) x 25. 168. Q.

patula. Biocal stn DS 98, x 75. 169-173, Q. laetifica. 3.02 x 3.90 mm. 172 (lasi rw) x 40. 173 x 70. 174. Q.

limatula, paratypc. Biocal stn CP 26. 4.52 x 6.90 mm. 175-178, Q. limalula. holotype. 3.60 x 5.40 mm. 177 (last
tw) x 40, 178 x 80.

Source : MNHN, Paris

86

BRUCE

MARSHALL

Quinnia limatula sp. nov.

Figs 174-178

Description. — Shell up to 6.90 mm wide,
broader than high, thin, umbilicate, spire 0.71-
1.11 x as high as aperture ; white, nacreous
through thin, translucent outer shell layer.

Protoconch 330 iim wide, tip minutely granu-
late, last half whorl smooth.

Teleoconch of up to 5.9 whorls ; lst whorl with
strong supramedian shoulder angulation, angu¬
lation weakening and vanishing over next half
whorl, subséquent whorls very weakly concave
or convex, periphery sharply angulate ; base
suddenly contracted, convex, flattened below
periphery. Axial riblets fine, sigmoidal, colla-
bral, entirely traversing lst 2 whorls, after
which obsolète in supramedian band, crisp on
lst 3.5 whorls, thereafter obsolète. Spiral threads
crisp, similar to axials on early whorls, multi-
plying by intercalation, becoming obsolète over
abapical half or two thirds of each whorl on 3rd
or 4th whorl, reappearing in subsutural zone on
last adult whorl. Base finely lirate over outer
third, umbilicus bounded by 3-5 smooth spiral
cords, innermost 3 most widely spaced and with
axial riblets in interspaces, médian third with
weakly defined grooves and obscure spiral fines.
Umbilicus deep, wall vertical, diameter 27.5-
29.6 % of shell diameter. Aperture subtrapezoi-
dal. Outer lip rim damaged, with narrow médian
tubular cavity in peripheral and basal angula¬
tion, from collabral growth fines retraction dcpth

3.7% and protraction depth 16.7% of shell
diameter ; basal notch concave ; peripheral notch
within peripheral angulation, not retracted.
Pariétal glazc thin. Inner lip (adult paratype)
rather thin, curved towards umbilicus, abapical
end broken but probably flexed to form a small
tooth.

Animal unknown.

Type data. Holotype mnhn (3.60 x
5.40 mm. 5.25 tw) : Bioc al, stn DW 79. Para-
type mnhn (4.52 x 6.90 mm, 5.9. tw) : Biocal
stn CP 26.

Distribution. OIT Ouvéa, Loyalty Islands
and Southern New Caledonia, 1320-1 740 m
(dead).

Remarks. Quinnia limatula differs from ail
known species of Quinnia in lacking a shoulder
angulation on ail but the earliest teleoconch
whorls. Unlike Q. palula, Q. cazioti and Q.
laetijica, this species probably has a tooth on the
aduit inner lip, a characler shared with possible
congeners Seguenzia ionica Watson, 1879 and S.
rushi Dali, 1927. Apart from the lack of a
shoulder angulation on later whorls, and the
probable presence of a tooth on the inner lip, Q .
limatula is essentially similar to typical Quinnia
species. The tubular cavities in the outer lip rim
at the peripheral and basal angulations are
characters hilherto unknown from this family.

Etymology. Somewhal polished (Latin).

Genus H AL YSTINA gen. nov.

Type species. — Halystina caledonica sp. nov.

Etymology. Diminutive of seguenziid genus
Halystes Marshall, 1988, which is an anagram of
the seguenziid genus Thelyssa Bayer.

Diagnosis. Shell 2.1 -4.0 mm high, depressed
or narrowly turbiniform, narrowly umbilicate.
Protoconch finely granulate, with or wilhout
2 fine spiral threads. Teleoconch whorls becom¬
ing almost flat-sided. Sculpture of collabral axial

and spiral riblets, axials well-developed through-
oul or soon becoming obsolète, with or without
stronger, angulating spirals on spire. Posterior
apertural notch broad, protraction dcpth 9.7-
25.2 % of shell diameter. Columella toothless.
External anatomy unknown (animais dried).
Radula similar to that in Halystes and Seguen¬
zia.

Remarks. Apart from the three species
described below, this genus contains Seguenzia
siberutensis Thiele, 1925 (off Siberut Island

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZ11DAE

87

Figs 179-193. Gcnus Halystina : 179. 180. 182. 183, Halystina vauhani. holotvpc. 1.80 x 1.70 mm. 182 x 55. 183
181, //. vauhani, paralype. “ Vauhan " sln 40, widih 1.80 mm. 184-i88, //. calédonien, holotype. 3.00 x
187 x 40, 188 x 85. 189-193, II. carinata, holotype. 3.40 x 3.18 mm. 192 x 33, 193 x 95.

x 110.—
2.70 mm.

Source : MNHN, Paris

88

BRUCE A. MARSHALL

western Sumatra, 750 m), and S. simplex (Bar-
nard, 1963) (ofT Cape Point, South Africa,

1 280 m). Il is thus équivalent to Seguenzia
Group III of Quinn (1983). Members of the
group are extremely similar to the abyssal type
species of Halystes Marshall, 1988 (H. chimaera
Marshall, 1988) in gross faciès, but dilTer collecti-
vely in attaining smallcr maximum size (height
2.1-4.0 mm, cf. 8.6 mm), and in having narrower
and deeper posterior notches (maximum depth
9.7-25.2 %, cf. 2.3 % of shell diameter). Although
Halystina is here interpreled as a discrète mono-
phyletic radiation, Halystes chimaera may prove
to be an aberrant member of the same group. in
which case Halystina might be better placed as a
subgenus. Until animais of Halystina species are
available for comparison with that of H. chi¬
maera, I prefer to treat Halystes and Halystina as
distinct, closely related généra. The central radu-
lar tooth in H. caledonica is shorter than in H.
chimaera. Both groups appear to be closely
related to Rotellenzia Quinn, 1987 (type species
Basilissa lampra Watson, 1879).

Halystina caledonica sp. nov.

Figs 184-188, 282, 283; Table 17

Description. Shell up to 3.35 mm high,
higher than broad, umbilicus mostly invaded by
inner lip, thin ; spire weakly cyrtoconoid, 0.77-
0.82 x as high as aperturc ; white, nacreous
through thin, translucent outer shell layer.

Table 17. — Halystina caledonica. Shell measurements (mm)
and countings. (Biocal, sln DS 04, DS 98).

Character

n

Range

Mean

SD

H

6

2.85-3.35

3.07

0.16

D

6

2.40-2.95

2.78

0.20

H/D

6

1.05-1.19

1.11

0.05

TW

6

4.40-4.50

4.45

0.05

Protoconch 330 u.m wide, minutely granulate.

Teleoconch of up to 4.5 whorls ; lst 2 whorls
convex, subséquent whorls fiat above rounded
periphery, base convex. Very minutely granulate
throughout. Axial riblets fine, crisp, sigmoidal,
collabral, traversing spire and base, evanescent
on outer part of umbilical wall. Spiral threads
fine, crisp, similar on spire and base, multiplying
by intercalation, numbering about 12 at start of

last adult whorl and 12-16 on base, an additional
thread on outer part of umbilical wall in most
adults. Umbilicus reduced to a shallow crescentic
dépréssion by invading inner lip. Aperture sub-
trapezoidal. Outer lip thin ; posterior notch
broad, retraction depth 6.9-7.0 % and protrac¬
tion depth 17.2-18.6% of shell diameter; basal
notch concave, apical rim slightly flared ; periph-
eral notch shallow, concave. Pariétal glaze thin.
Inner lip almost straight, toothless.

Animal unknown (dried).

Radula (Figs 282, 283). Central tooth sub-
ovate, longer than broad, thin in section, flexible,
cutting area curving forward from shaft, nar-
rowly angulate, terminal cusp large, accessory
cusps fine, about 8 on each side, shaft tapered to
horizontal basal plate. Latéral teeth broad, thin
in section, flexible, cutting area narrowly angu¬
late, terminal cusp large, 4-6 fine, accessory cusps
on each side. Marginal teeth slender, terminal
cusp of each very long and slender, a few fine
accessory cusps on outer edge.

Jaw not found, reduced or absent.

Type data. Holotype (3.00 x 2.70 mm,
4.5 tw) and paratype mnhn ; Biocal, stn DS 04.
Paratypes (3 mnhn, I nmnz) : Biocal, stn DS 98.

Distribution. Between New Caledonia and
Lifou, Loyalty Islands, 2 340-2 470 m, living at
2 340 m.

Remarks. Compared with Halystina siheru-
tensis, which it much resembles in general faciès,
H. caledonica difîers in sculptural details, in
attaining maturity at larger size (height 3.40-
4.00 mm cf. 2.40 mm) and in having a larger
protoconch (width 330 ^m cf. 230 nm).

Etymology. (New) Caledonian.

Halystina carinata sp. nov.

Figs 189-193; Table 18

Description. Shell up to 4.00 mm high,
slightly higher than broad at maturity, thin,
umbilicus invaded by inner lip, sometimes with a
fine crescentic chink, spire 1.13-1.15 x as high as
aperture ; white, nacreous through thin, translu¬
cent outer shell layer.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

89

Protoconch 300-310 jxm wide, minutely granu-
late.

Teleoconch of up to 5.75 whorls ; shoulder
angulated by sharp-edged keel, ramp and side
more or less fiat ; periphery rounded, with
smooth angulate keel. base convex. Shoulder keel
strongly supramedian on lst whorl, descending
to submedian position, smooth ; subsutural angu¬
lation commencing on about mid 3rd whorl.
summit undulant throughout. Axial riblets colla-
bral, fine, crisp on lst 2 whorls, weakening on
3rd whorl, becoming obsolète on 4th whorl,
sometimes reappearing on ramp on last part
of last adult whorl. Spiral threads fine, crisp,
multiplying by intercalation, commencing on
3rd whorl, covering last adult whorl from suture
to periphery and between periphery and outmost
basal spiral. Base with 8-10 smooth, subequal
spiral cords, interspaces considerably wider than
each spiral. Aperture subtrapezoidal. Outer lip
thin, slightly thickened within ; posterior notch
retraction depth 9.6-10.7 % and protraction depth
21.5-25.2 % of shell diameter ; basal notch con¬
cave, gently flared ; peripheral notch small, shal-
low. Pariétal glaze thin. Inner lip rather thick,
spreading into and almost or entirely infilling
umbilicus, almost straight, toothless.

Animal unknown.

Table 18. Halysiina carinata. Shell measurements (mm)
and countings. (Biocal, sln DW 79).

Character n

Range

Mean

SD

H 8

3.35-4.00

3.54

0.20

D 8

3.03-3.38

3.22

0.12

H/D 8

1.05-1.18

1.09

0.04

TW 8

5.20-S.75

5.36

0.19

Type data. Holotype mnhn (3.40 x
3.18 mm, 5.4 tw) and 11 paratypes (ams, bmhn,
mnhn, nmnz, nmp, usnm) : Biocal, stn DW 79.

Distribution. Off Ouvêa, Loyalty Islands,
1 320-1 380 m (dead).

Remarks. Halystina carinata dififers markedly
from its congeners in having angulate subsutural,
shoulder and peripheral keels. The South Afri-
can H. simplex (Barnard, 1963) resembles it in
having a shoulder angulation, but in H. carinata
the angulation is set lower on later whorls, the
spire is more narrowly conical, there are more

numerous basal spire cords, while the umbilicus
is closed instead of fully open.

Etymology. Keeied (Latin).

Halystina vaubani sp. nov.

Figs 179-183; Table 19

Description. — Shell up to 2.10 mm high,
about as high as broad, thin, narrowly umbilicate,
spire 0.75-0.92 x as high as aperture ; white,
nacreous through thin, translucent outer shell
layer.

Table 19. Halystina vaubani. Shell measurements (mm)
and countings. (" Vauban ”, stn 40).

Character

n

Range

Mean

SD

H

10

1.72-2.10

1.92

0.12

D

10

1.70-1.82

1.78

0.04

H/D

10

0.98-1.15

1.07

0.06

TW

10

3.80-4.40

4.15

0.17

Protoconch 330-400 |xm wide, sculptured with
minute granules and 2 fine spiral threads.

Teleoconch of up to 5.75 whorls; lst spire
whorl convex, subséquent whorls almost fiat,
with low angulate keels at about adapical third
and subsuturally ; periphery broadly angulate ;
base convex, on last adult whorl becoming
shallowly concave from midway between periph¬
ery and umbilicus to innermost 2 spiral threads.
Last part of last adult whorl becoming steeper-
sided. Axial riblets fine, sigmoidal, collabral,
crisp on spire, weaker on base where typically
absent from a médian spiral band. Spiral threads
fine, crisp, traversing axials, multiplying by inter¬
calation, numbering 7-9 on spire at start of last
whorl, and 10-12 on base. Very minutely granu-
late throughout. Umbilicus conical, diameter
18.3-23.1 % of adult shell diameter. Aperture
subquadrate. Outer lip thin ; posterior notch
broad, shallow, rim damaged, from collabral
sculpture rétraction depth 5.8 % and protraction
depth 9.7 % of shell diameter ; basal notch
concave, rim gently flared ; peripheral notch
shallow, angulate. Pariétal glaze thin. Inner lip
thin, almost straight, toothless.

Animal unknown.

Source : MNHN, Paris

90

BRUCE A. MARSHALL

Type data. Holotype mnhn (1.80 x
1.70 mm, 4.0 tw) and 32 paratypes (ams. bmnh,
mnhn, nmnz, nmp, usnm) : “ Vauban ", stn 40.

Distribution. — Off Southern New Caledo-
nia, 250-350 m (dead).

Remarks. — Compared with H. siberutensis
(Figs 136-138) to which it is closely similar, H.
vaubani differs in having considerably narrower,
more crisply defined spiral threads on the lasl
adult whorl, and in having a low angulation at
the adapical third on the spire whorls, while the
innermost two basal spiral threads are finer, and

the inner base is more shallowly concave at the
same stage of growth. Moreover, H. vaubani
appears to attain maturity at smaller size (maxi¬
mum height 2.10 mm, cf. 2.40 mm), and does not
become as distinctly pupoidal in outline at
maturity. Of the 14 syntypes of H. siberutensis
(Zoological Muséum, Berlin) the specimens here
chosen as lectotype (Figs 136-138) closely matches
the original illustration (Thiele, 1925, PI. 1,
figs 13-14) and agréés well with the original given
dimensions (2.4 x 1.9 mm).

Etymology. Named after N. O. "Vau¬
ban ".

Genus SEGUENZIA Jeffreys, 1876

Seguenzia Jeffreys, 1876 : 200. Type species (by
monotypy) : Seguenzia formosa Jeffreys, 1876 ;
Recent, North Atlantic.

Seguenzia chelina Marshall, 1983
Figs 194-197 ; Table 20

Seguenzia chelina Marshall, 1983 : 240, figs 2k-o.

Type data. Holotype nzoi h. 371 : P 929,
40°42.8' S, 167°50.0' E, E slope of Tasman Basin,
off Westport, New Zealand, 1 029 m, 18 April
1980, R. V. "Tangaroa".

Other material examined. New Caledonia and
Loyalty Islands (5 specimens) — Biocal stn CP 75 (1
mnhn)! — Stn DW 80 (3 mnhn. 1 nmnz). New
Zealand (5 specimens) : BS 846 (O 592), 37 l, 04.3' S,
176”26.6'E, SE of Aldermen Is, dead, 807-872 m,
23 January 1981, R. V. “ Tangaroa " (3 nmnz) ; P 942,
41°00.6' S, 169°06.0' E, E slope of Tasman Basin, off
Westport. dead, 914 m, 24 April 1980, R. V. “ Tan¬
garoa " (2 nzoi).

Table 20. — Seguenzia chelina . Shell measurements (mm)
and countings.

H

D

H/D

TW

1.22

5.60

Biocal, stn

DW 80

1.22

5.90

Biocal, stn

DW 80

2.61

1.15

6.00

Biocal, stn

DW 80

2.60

1.15

6.25

Holotype

2.50

1.22

6.00

BS 846

2.95

1.12

6.00

BS 846

2.80

1.18

6.00

P 942

3.35

2.65

1.26

5.80

BS 846

Distribution. New Caledonia and Loyalty
Islands (825-980 m), and New Zealand (807-
1 029 m).

Remarks. Specimens from off New Caledo¬
nia and the Loyalty Islands differ from the
holotype and additional material from the Tasman
Basin in having the umbilicus only partly invaded
by the inner lip and thus more open, but are
otherwise indistinguishable. Two of the three
New Zealand specimens laken off the Aldermen
Islands hâve the umbilicus inlermediate in width
between the extremes, so there would seem to be
north-south clinal intergradation in umbilicus
width. It transpires that the holotype is atypical
in lacking spiral cords from a broad zone beside
the umbilical chink. Ail other New Zealand
specimens hâve spiral cords that extend to the
umbilical rim (Fig. 197), numbering 7-9 in New
Zealand material, and 8 or 9 in specimens from
New Caledonia and the Loyalty Islands. See
Discussion page 107.

Seguenzia praeceps sp. nov.

Figs 198-202; Table 21

Description. Shell up to 4.55 mm high,
higher than broad. thin, anomphalous, spire
1.19-1.35 x as high as aperture ; white, nacreous
through thin, translucent outer shell layer.

Protoconch 300-330 p.m wide, minutely granu-
late.

Teleoconch of up to 6.6 whorls, shoulder and

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SF.GUENZIIDAE

91

Figs 194-207. — Genus Seguenzia : 194-196, Seguenzia chelina. Biocal stn DW 80. 3.15 x 2.65 mm. 197, S. chelina, ofT
Aldermen Is, New Zcaland, 807-872 m (nmnz m. 95411), width 2.70 mm. 198-202, S. praeceps, holotype,
4.15 x 3.25 mm. 201 x 30. 202 x 85. 203-207, S. metivieri , holotype, 2.88 x 2.52 mm. 206 x 45. 207 x 90.

Source : MNHN, Paris

92

BRUCE A. MARSHALL

periphery angulated by sharp-edged keels, ramp
shallowly concave, concave between periphery
and outermost basal spiral, most deeply concave
between shoulder and periphery ; base gently
contracted, convex. Very minutely granulate
throughout. Shoulder keel at adapical third, its
summit on last whorl with distinct bevel bounded
by spiral threads ; a finely serrate subsutural
angulation commencing late on 4th whorl. Axial
riblets fine, sigmoidal, crisp on spire and base to
innermost basal spiral, numbering 13-18 per mm
between spire keels at end of 5th whorl. spiral
threads finer than axials, multiplying by interca¬
lation, covering spire and periphery to outermost
basal spiral. Base with 7-9 similar, prominent
spiral cords, interspaces considerably broader
than each spiral. Aperture subrhomboidal. Outer
lip thin, posterior notch deep, retraction depth
14.0-18.4% and protraction depth 53.2-58.4%
of shell diameter, apical rim flared ; basal notch
U-shaped, apical rim flared ; peripheral notch
angulate, well retracted. Pariétal glaze thin. Inner
lip concave, sharply flexed at base to form
prominent tooth, broadly channelled below.

Animal unknown.

Table 21. Seguenzia praeceps. Shell measurements (mm)
and countings. (Biocal, stn DW 79).

Character

n

Range

Mean

SD

H

9

4.00-4.55

4.22

0.22

D

9

3.15-3.35

3.23

0.08

H/D

9

1.25-1.38

1.30

0.04

TW

9

6.10-6.60

6.41

0.18

Type data. — Holotype mnhn (4.15 x
3.25 mm, 6.10 tw) and 11 paratypes (ams, bmnh,
MNHN, NMNZ, nmp, usnm) : Biocal, stn DW 79.

Other material examined (3 specimens mnhn). —
Biocal, stn DS 59 (2). — Stn DS 98 (1).

Distribution. — Off Ouvéa, Loyalty Islands
and Southern New Caledonia, 1 320-2 650 m
(dead).

Remarks. — Among previously described
species, 5. praeceps seems closest to S. fulgida
Marshall, 1983, based on two specimens taken at
1 760-1 799 m off Westland, New Zealand. The
New Caledonian species differs primarily in having
more crisply-defined axial riblets on later whorls.

including the base, in having doser shoulder
and peripheral keels, and in attaining maturity
at smaller size, S. fulgida attaining 6.10 mm
in height. It also closely resembles syntypes
(usnm 181650) and nmnz specimens of S. for-
mosa Jeffreys, 1876 (off Portugal, 4 220-4 380 m)
and the holotype (nsmt) of 5. mirabilis Okutani,
1964 (off Japan, 3 150-3 550 m) in general faciès]
differing from S. formosa in having a smaller
protoconch (diameter 300-330 nm, cf. 370 j^m)
and a sharper subsutural angulation, and from
S. mirabilis in being smaller relative to the
number of whorls and in having considerably
weaker spiral threads on the spire. The unde-
scribed Philippine species illustrated by Quinn
(1983b, fig. I) is superficially similar, but differs
in having a much stronger tooth on the inner lip.

Etymology. Steep (Latin).

Seguenzia metivieri sp. nov.

Figs 203-207

Description. Shell up to 2.88 mm high,
slightly higher than broad, thin, anomphalous,
or with narrow umbilical chink, spire 1.15 x as
high as aperture ; white. nacreous through thin,
translucent outer shell layer.

Protoconch 280 [xm wide, finely granulate.

Teleoconch of up to 5.80 whorls, shoulder and
periphery angulated by sharp-edged keels, ramp
almost fiat, side concave, shallowly concave
between periphery and outermost basal spiral,
base rather gently contracted, convex. Very
minutely granulate throughout. Shoulder angu¬
lation at about adapical third, a low angulation
bordering suture on last adult whorl. Axial
riblets fine, sigmoidal. crisp on spire and on base
to outermost basal spiral, obsolète on adapical
side of shoulder keel, obsolète elsewhere on base,
numbering 17 per mm between keels at end of
5th whorl. No spiral threads. Base with 8 spiral
cords, outermost cord angulate in section, most
prominent, others considerably lower, interspaces
similar, considerably broader than each spiral.
Umbilicus narrow, either fully invaded by inner
lip, or partially invaded to form an elliptical
chink. Aperture subrhomboidal. Outer lip thin,
posterior notch deep, retraction depth 10.2 %
and protraction depth at least 24 % shell diame¬
ter (tip of labial projection broken), apex flared ;

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

93

basal notch shallow, concave, apex slightly fla-
red ; peripheral notch smallest, roundly angulate.
Pariétal glaze thin. Inner lip thick, deeply curved
into umbilicus, sharply flexed at base to form
strong tooth, channelled below.

Animal unknown.

Type data. — Holotype (2.88 x 2.52 mm,
5.50 TW) and paratype (2.75 x 2.45 mm, 5.80 rwj
mnhn : Biocal, stn DW 80.

Distribution. — Off Ouvea, Loyalty Islands,
900-980 m (dead).

Remarks. Seguenzia metivieri bears a close
superficial resemblance to the New Zealand
species S. transenna Marshall, 1973, from which
il difiers primarily in lacking spiral threads on
the teleoconch, and in having the axial riblets
weaker between the periphery and outermost
basal spiral, and obsolète elsewhere on the base,
while the tooth on the inner lip is much stronger.
Moreover, the axial riblets on the shoulder are
shorter. extending almost to the summit of the
keel in S. transenna. S. metivieri occurred toge-
ther with S. chelina at the type locality.

Etymology. Named after Bernard Meti-
vier (mnhn) who participated on the Biocal
campaign.

Seguenzia richeri sp. nov.

Figs 208-212

Description. Shell (holotype) 3.55 mm
high, slightly higher than broad, thin, umbilicate,
spire as high as aperture ; white, nacreous through
thin, translucent outer shell layer.

Protoconch 370 ^m wide, minutely granulate.

Teleoconch of 5.0 whorls ; shoulder and periph¬
ery angulated by sharp-edged keels, ramp and
side concave, concave between periphery and
outermost basal spiral ; base evenly contracted,
convex. Very minutely granulate throughout.
Shoulder keel at about adapical third, adapical
edge distinctly bevelled on last 2 whorls, bevel
bounded by crisp spiral thread ; a low, finely
serrated subsutural angulation on last whorl.
Axial riblets fine, crisp, sigmoidal, traversing
spire and base onto outer part of umbilical wall.
numbering 13 per mm on spire between keels at

end of 5th whorl. Spiral threads crisp, similar to
axials, multiplying by intercalation, covering
spire and periphery to outermost basal spiral.
Base with 6 similar prominent spiral cords ;
interspaces similar, considerably broader than
each spiral. Umbilicus deep, rim angulate, dia-
meter 19.3 % of shell diameter. Aperture sub-
rhomboidal. Outer lip thin ; posterior notch
deep, retraction depth 6.45 % of shell diameter,
protraction depth unknown (rim damaged) but
at least 34 % of shell diameter ; basal notch well
developed, flared ; peripheral notch in angula¬
tion, slightly retracted. Pariétal glaze very thin.
Inner lip rather thin, rolled outwards at rim,
deeply curved into umbilicus, sharply flexed at
base to form prominent tooth, narrowly channel¬
led below.

Animal unknown.

Type data. Holotype mnhn (3.55 x
3.10 mm, 5 tw) : Biocal, stn CP 23.

Distribution. — Off Southern New Caledo-
nia, 2 040 m (dead).

Remarks. - Seguenzia richeri closely resem-
bles S. metivieri sp. nov.. and the New Zealand
species S. transenna Marshall, 1973. in shape,
diflering from both in having a larger proto¬
conch and a wider, open umbilicus. U differs
further from 5. metivieri in lacking the bevelled
edge on the shoulder keel, in having strong axial
sculpture on the base, and in having fewer (6 cf.
8), more widely spaced cords on the base, while
S. transenna has finer basal sculpture with 11 or
12 spiral cords, and a much weaker tooth on the
inner lip.

Etymology. — Named after Bertrand Richer
de Forges, who participated on the Biocal
campaign.

Seguenzia emmeles sp. nov.

Figs 213-217

Description. Shell (subadult holotype)
3.55 mm high, higher than broad, thin, umbili¬
cate, spire 1.25 x as high as aperture; white,
nacreous through thin, translucent outer shell
layer.

Protoconch (holotype and paratype) 330 ,um
wide, tip bulbous.

Source : MNHN, Paris

94

BRUCE A. MARSHALL

Figs 208-222 Genus Seguenziu : 208-212, Seguenzia richeri, holotype, 3.55 x 3.10 mm, 211 x 35, 212 x 85
emmeles, holotype, 3.55 x 2.90 mm, 2l6 x 30. 217, S. emmeles , paralype, Biocal stn 59. x 70

eutvches, holotype, 2.90 x 3.00 mm, 221 x 40, 222 x 80.

213-216, S.
218-222, S.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZI1DAE

95

Teleoconch (holotype) of 5.6 whorls ; shoulder
and periphery angulated by similar, sharp edged
keels, ramp and side concave, concave between
periphery and outermosl basal spiral ; base evenly
contracted, convex. Shoulder keel at about ada-
pical third, a low, finely serrate subsutural
angulation commencing on 3rd whorl. Axials fine,
crisp, sigmoidal, collabral, entirely traversing
spire and base, evanescent deep within umbilicus,
numbering 12 per mm on spire between keels at
end of 5th whorl. Spiral threads crisp, numerous,
similar, multiplying by intercalation, covering
spire, periphery and base to outermost spiral, a
few threads in each interspace of inner basal
spirals. Base with 4 strong spiral cords, outer¬
most angulate in section, most prominent, others
lower and similar, interspaces much wider than
each spiral. Umbilicus deep, diameter 25.3 % of
shell diameter. Aperture subtrapezoidal. Outer
lip thin, retraction depth of posterior notch 6.9 %
of shell diameter, protraction depth unknown
(rim damaged) ; basal notch concave ; peripheral
notch small, angulate, slightly retracted. Pariétal
glaze very thin. Inner lip thin, shallowly curved
towards umbilicus, weakly flexed at base, tooth-
less.

Animal unknown.

Type data. Holotype (3.55 x 2.90 mm,
5.6 tw) and immature paratype mnhn : Biocal,
stn DS 59.

Other material examined (3 immature specimens
MNHN). — Biocal, stn DS 04 (2). Stn CP 72 (1).

Distribution. OfT Southern New Caledo-
nia, 2 100-2 650 m (dead).

Remarks. — Compared with S. richeri to
which it has a general resemblance, S. emmeles
differs in having a more excert protoconch,
a more narrowly conical spire, and in lacking
the clearly defined bevel on the adapical side of
the summit of the shoulder keel. 5. mirabilis
Okutani, 1964 (ofT Japan, 3 150-3 350 m; holo¬
type at Geological Institute, University of Tokyo,
rm 8811) is even doser in general faciès, but
differs in sculptural details, in having a narrow
umbilical chink, and in having 7 instead of
4 spiral cords on the base.

Etymology. — Harmonious (Greek).

Seguenzia eutyches sp. nov.

Figs 218-222

Description. — Shell up to 3.40 mm high.
slightly broader than high, thin, umbilicate, spire
0.78-1.03 x as high as aperture ; white. nacreous
through thin, translucent outer shell layer.

Protoconch 300jim wide, minutely granulate.

Teleoconch of up to 5.2 whorls ; shoulder and
periphery angulated by sharp-edged keels, ramp
and side concave, concave between periphery and
outermost basal spiral ; base gently contracted,
weakly convex. Very minutely granulate through-
out. Shoulder keel at adapical third, similar to
peripheral keel ; low, finely serrate subsutural
angulation commencing on 2nd whorl. Axial
riblets fine, crisp, sigmoidal, strong over spire to
outermost basal spiral, finer on inner base,
evanescent on outer part of umbilical wall,
numbering 15 per mm at end of 5th whorl. Spiral
threads almost obsolète, présent on spire and
between periphery and outermost basal spiral.
Base with 8 crisp spiral cords, outermost angulate
in section, most prominent, next spiral finest, inner
spirals low and and similar. spiral interspaces
considerably wider than each spiral. Umbilicus
deep, diameter 21.6-27.7% of shell diameter.
Aperture subrhomboidal. Outer lip thin. rim
damaged, basal notch concave, apical rim slightly
flared. Pariétal glaze thin. Inner lip thick, rim
tightly folded towards umbilicus, gently curving
towards umbilicus, gently flexed at base to form
small rounded looth, shallowly channelled below.

Animal unknown.

Type data. Holotype (2.90 x 3.00 mm,
5.2 tw) and paratype (3.40 x 3.55 mm, 5.1 tw)
mnhn : Biocal, stn DW 70.

Distribution. — Off Southern New Caledo-
nia, 965 m (dead).

Remarks. — Seguenzia eutyches most closely
resembles S. richeri sp. nov. but differs in a
number of characters, including the very much
weaker spiral threads, the earlier appearance of
the subsutural angulation, the more shallowly
concave inner lip, and the considerably weaker
tooth. It differs further in the lack of a thread-
bounded bevel on the shoulder keel, and in the

Source : MNHN, Paris

96

BRUCE A. MARSHALL

straighter axial riblets on the ramp on the early
teleoconch whorls.

Etymology. — Good luck (Greek).

Seguenzia wareni sp. nov.

Figs 223-227 ; Table 22

Description. Shell up to 2.90 mm high,
slightly higher than broad, thin, narrowly umbi-
licate. spire 0.97-1.07 x as high as aperture ;
white, nacreous through thin, translucent outer
shell layer.

Proioconch 270-300 |im (mostly 270 |xm) wide,
finely granulate.

Table 22. Seguenzia wareni. Shell measurcmcnts (mm)
and countings. (Biocal, sln DW 79).

Character

n

Range

Mean

SD

H

10

2.55-2.90

2.69

0.11

D

10

2.30-2.55

2.45

0.07

H/D

10

1.04-1.15

1.09

0.04

TW

10

4.80-5.20

4.94

0.12

UD%

10

16.8-20.8

18.9

1.14

Teleoconch of up to 5.2 whorls ; shoulder and
periphery angulated by sharp-edged keels, ramp
and side concave, concave between periphery
and outermost basal spiral ; base evenly con-
tracted, convex. Very minutely granulate through-
out. Shoulder keel strongly supramedian at first,
descending to adapical third. Axial riblets, fine,
crisp, sigmoidal, entirely traversing spire and
base to umbilical rim, numbering about 33 per
mm between spire keels at end of 5th whorl.
Spiral threads similar, similar to axials, mul-
tiplying by intercalation, extending from suture
to periphery, a few developing on last adult
whorl between periphery and outermost basal
spiral. Base with 7 similar, prominent spiral
cords, interspaces considerably broader than
each spiral. Frequently a weaker cord immedia-
tely within umbilicus. Umbilicus deep, rim angu-
late, diameter 16.8-20.8 % of adult shell diame-
ter. Aperture subrhomboidal. Outer lip thin ;
posterior notch deep, retraction depth 12.9-
17.3% and protraction depth 54.8-55.4% of
shell diameter, apical rim flared ; peripheral and
basal notches similar, U-shaped, flared. Pariétal

glaze thin. Inner lip strongly curved towards
umbilicus, sharply flexed at base to form promi¬
nent tooth, channelled below.

Animal unknown.

Type data. Holotype mnhn (2.80 x
2.45 mm, 5 tw) and 20 paratypes (ams, bmnh,
MNHN, NMNZ, NMP, USM) : BlOCAL, Stn DW 79.

Distribution. OIT Ouvéa, Loyalty Islands,
1 320-1 380 m (dead).

Remarks. Compared with S. richeri, which
it much resembles in shape and in having a
narrow, open umbilicus, S. wareni diflers in
being smaller with a smaller protoconch, and in
having finer and doser reticulate sculpture.

The unusually deep, U-shaped peripheral notch
in the mature outer lip is an exlremely distinctive
character. S. wareni occurred together with 5.
chariessa at the type locality.

Etymology. - This species is named after
Anders Warén (Swedish Muséum of Natural
History, Stockholm), who provided critical com¬
parative material and who designed the excellent
rock dredge with which most of the présent
material was obtained.

Seguenzia malara Marshall, 1988
Figs 228-232

Seguenzia matara Marshall, 1988 : 242, figs 2 k-o, 4 g.

Type data. Holotype nzoi h. 453 : Q 696,
42 0 36.1'S, 169°34.8' E, ofT Westland, New Zea-
land. alive, 935-920 m, 21 February 1982, R. V.
“ Tangaroa ".

Other material examined (2 spécimens mnhn). —
Biocal, stn DW 80.

Distribution. OIT Ouvéa, Loyalty Islands
(dead, 900-980 m), and New Zealand (alive, 750-
1 029 m).

Remarks. The two New Caledonian spéci¬
mens fall within the range of variation in shell
morphology exhibited by New Zealand type
material of S. malara, and would thus seem to be

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

97

Figs 223-237. Gcnus Seguenzia : 223, 225-227, Seguenzia wareni. hololype. 2.80 x 2.45 mm. 226 * 45, 227 x 95. 224,

.V. wareni, paratype, Biocai. sln DW 79, x 20. 228-232, S. malara. Biocal stn DW 80. 3.20 x 3.40 mm. 231 x 45

232 x 80. 233-237, 5. eidalima. hololype, 3.27 x 3.10 mm. 236 x 45. 237 x HO.

Source : MNHN, Paris

98

BRUCE A. MARSHALL

conspecific. S. malara is similar to S. elegans
Jeffreys, 1885 and S. nipponica Okutani, 1964 in
general faciès. It differs from both in that the
outermost basal spiral and peripheral keel are set
further apart. It differs further from 5. elegans in
attaining maturity of larger size and in having a
wider umbilicus, while the last adult whorl is
more strongly convex with more prominent
shoulder and peripheral keels. When introducting
5. malara I did not indicate its affinities with S.
nipponica because New Zealand specimens see-
med to differ widely from the original illustration
of the holotype (Okutani, 1964. pl. 6, fig. 1.
cf. Marshall. 1988. fig. 2 k). Comparisons of
the type material, however, reveal that they are
in fact closely related. Of the three species,
S. nipponica and S. elegans are the most similar,
5. nipponica differing from S. elegans in attaining
larger size and in having fewer, more widely spaced
axial riblets on the spire. A closely related species
is described below. See Discussion page 107.

Seguenzia eidalima sp. nov.

Figs 233-237

Description. Shell (holotype) 3.27 mm
high, about as broad as high, thin, umbilicate,
spire 0.80 x as high as aperture ; white, nacreous
through thin, translucent outer shell layer.

Protoconch 270 [im wide, surface mostly etched
away, remaining surface granulate.

Teleoconch of 5.5 whorls, shoulder and periph-
ery angulated by sharp-edged keels, ramp and
side shallowly concave, shallowly concave between
peripheral and outermost basal spiral, base evenly
contracted, weakly convex. Very minutely granu¬
late throughout. Shoulder keel near adapical
third, rather low, peripheral keel stronger ; low,
finely serrate subsutural angulation commencing
on 2nd whorl. Axial riblets fine, crisp on spire
and on outer half of base, weaker on inner half
of base, numbering 10 per mm between keels at
end of 5th whorl. Spiral threads multiplying by
intercalation, crisp, that surmounting subsutural
angulation becoming as strong as axials, others
finer and similar, covering spire and occupying
spaces between periphery and outermost 2 basal
spirals. Base with 13 crisp spiral cords, inter-
spaces considerably wider than each spiral, outer
2 spirals angulate in section, most prominent,
others considerably lower. Umbilicus deep, rim

narrowly rounded and overhanging wall that is
sigmoidal in section, diameter 26.7 % of shell
diameter. Aperture subrhomboidal. Outer lip
thin, posterior notch apex flared, retraction
depth 10.7 % of shell diameter, protraction
depth unknown (labial projection broken) ; basal
notch concave, no peripheral notch. Pariétal
glaze thin. Inner lip rather thick, tightly rolled
outwards at rim, deeply curved towards umbili¬
cus, strongly flexed at base to form very promi¬
nent tooth, deeply channelled below.

Animal unknown.

Type data. Holotype mnhn (3.27 x

3.10 mm, 5.5 tw) : Biocal, stn CP 26.

Distribution. Off Southern New Caledo-

nia, 1 618-1 740 m (dead).

Remarks. Seguenzia eidalima closely resem-
bles type material of the North Atlantic species
5. elegans Jcffrcys, 1885 (syntypes bmnh 885.11.5
2628-30, 2587-88) and the Japanese S. nipponica
Okutani, 1964 (holotype Department of Geo-
logy, University of Tokyo rm 8812; paratype
nsmt Mo 64682). It differs from S. nipponica in
being smaller relative to the number of whorls
(S. nipponica paratype 4.20 x 4.05 mm, 4.5 tw),
and from S. elegans in having a wider umbilicus
(diameter 26.7 % of shell diameter, cf. 19.7 %).
It differs further from both in that the outer part
of the umbilical wall is concave and overhung by
the umbilical rim.

Etymology. Shapely (Greek).

Seguenzia iota sp. nov.

Figs 238-242 ; Table 23

Description. Shell up to 2.60 mm high,

slightly higher than broad, thin, narrowly umbi¬
licate, spire 0.95-1.09 x as high as aperture;
white, nacreous through thin, translucent outer
shell layer.

Protoconch 230-270 fim wide, coarsely granu¬
late throughout and with 2 spiral threads.

Teleoconch of up to 5.75 whorls ; shoulder and
periphery strongly angulated by prominent,
sharp-edged keels, ramp shallowly concave, shal¬
lowly concave between periphery and outermost
basal spiral, more deeply concave between shoul-

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZ1IDAE

99

Figs 238-252. Genus Seguenzia : 238, 239, Seguenzia iota, paratypcs. " Vaubun" sln 40. 2.30 x 2.10 (238). 2.05 :

240-242, S. iota, holotype, 2.23 x 2.01 mm. 241 x 60. 242 x 110. - 243, 244, 246, 247, S. cha
holotypc. 5.30 x 4.30 mm. 246 x 25. 247 x 70. 245, 5. chariessa , paratype. Biocal stn CP 75. width 3.95 n

248-251, S. levii, holotype, 5.00 x 3.55 mm. 251 x 25. 252, 5. levii. paratype. Biocai. stn CP 72. x 80

Source : MNHN, Paris

100

BRUCE A. MARSHALL

der keel and periphery ; base gently contracted,
convex. Very minutely granulate throughout.
Shoulder keel at about adapical third, summit
slightly upturned. Axial riblets fine, sigmoidal,
crisp on spire and base to outermost basal spiral,
developing between outermost 2 basal spirals on
last adult whorl, obsolète elsewhere on base,
numbering 16-22 per mm between keels at end of
5th whorl. Spiral threads crisp, numerous, multi-
plying by intercalation, covering spire, devel¬
oping between periphery and outermost basal
spiral on last adult whorl, and later, frequently
between outermost 2 basal spirals. Base with 5 or
6 crisp, similar spiral cords, interspaces consid-
erably wider than each spiral. Umbilicus deep,
outer part of wall shallowly convex, concave
behind inner lip, diameter 12.3-17.7% of adult
shell diameter. Aperture subrhomboidal. Outer
lip thin, posterior notch deep, apical rim strongly
flared radially and adapically, retraction depth
21.6-27.0% and protraction depth 65-73 % of
shell diameter ; peripheral notch only slightly
retracted though strongly radially flared at rim ;
basal notch concave, prominently flared. Pariétal
glaze thin. Inner lip rather thick, rim tightly
folded towards umbilicus, deeply curved towards
umbilicus, sharply flexed at base to form strong
tooth, channelled below.

Animal unknown.

Table 23. — Seguenzia iota. Shell measuremcnts (mm) and
countings. (“ Vauban ", stn 40).

Character

n

Range

Mean

SD

H

11

2.00-2.60

2.31

0.18

D

11

1.88-2.12

1.97

0.07

H/D

11

1.06-1.30

1.17

0.07

TW

11

4.90-5.75

5.31

0.28

IJD%

11

12.1-17.7

14.7

2.03

Type data. Holotype (2.23 x 2.01 mm,
5 tw) mnhn and 679 paratypes (ams, bmnh,
mnhn, nmnz, nmp, usm) : “ Vauban ”, stn 40.

Distribution. — Off southem New Caledo-
nia, 250-350 m (dead).

Remarks. Among previously described

species of Seguenzia. S. iota is rendered highly
distinctive by its small size and very strongly
developed apertural features.

Etymology. Very small (Greek).

Seguenzia chariessa sp. nov.

Figs 243-247 ; Table 24

Description. — Shell up to 5.75 mm high,
narrowly conical, considerably higher than broad,
thin, anomphalous, spire 1.32-1.36 x as high as
• aperture ; white, nacreous through thin, translu-
cent outer shell layer.

Proloconch 330-350 jxm wide (mostly 330 pm),
minutely granulate.

Teleoconch of up to 7.3 whorls ; shoulder and
periphery strongly angulated by smooth, sharp-
edged keels, ramp, side and basal spiral inter¬
spaces shallowly concave ; base evenly contrac¬
ted, convex. Shoulder keel strongly supramedian
at first, descending to almost médian position,
summit slightly upturned ; subsutural angulation
strong, smooth, commencing on 2nd whorl.
Axial riblets fine, sigmoidal. crisp on early
whorls, less crisply defined on later whorls,
extending over spire and base to umbilical rim,
numbering 13 per mm between shoulder keel and
periphery at end of 5th whorl. Spiral threads
absent. Base with 6 or 7 strong, smooth spiral
cords, interspaces considerably broader than
each spiral. Aperture subrhomboidal. Outer lip
thin ; posterior notch deep, retraction depth
12.1-14.6 % and protraction depth up to at least
61 % of shell diameter (tip of labial projection
broken), apical rim flared ; basal notch U-shaped,
rim flared ; peripheral notch shallow, concave,
flared. Pariétal glaze thin. Inner lip thick, deeply
curved away from aperture, strongly flexed at
base to form strong, narrowly angulate tooth,
channelled below.

Animal unknown.

Table 24. Seguenzia chariessa. Shell measurcments (mm)
and countings. (Biocal, stn CP 75, DW 80).

Character

«

Range

Mean

SD

H

7

4.70-5.75

5.30

D

7

3.65-4.25

3.92

0.20

H/D

7

1.21-1.41

1.35

0.05

TW

7

6.50-7.30

6.90

0.26

Type data. — Holotype (5.30 x 4.30 mm,
6.75 tw) mnhn, and 81 paratypes (ams, bmnh!
MNHN, NMP, NMNZ, USM) : BlOCAL, Stn CP 75.
Paratypes (12 mnhn) : Bioc:al , stn DW 79 (2)

Stn DW 80 (10).

Source : MNHN, Paris

MOLLUSCA OASTROPODA : SEGUENZIIDAE

101

Distribution. — OfT Ouvéa, Loyalty Islands
and southem New Caledonia, 825-1 380 m, living
at 825-860 m.

Remarks. Seguenzia chariessa is rendered
very distinctive by the combination of large size,
tall spire, lack of spiral threads, and rather weak
axial riblets.

Etymology. Graceful (Greeck).

Seguenzia levii sp. nov.

Figs 248-252

Description. — Shell up to 5.00 mm high,
narrowly conical, considerably higher than broad,
thin, becoming rather thick, very narrowly umbi-
licate, spire 1.58 x as high as aperture ; white.
nacreous through thin, translucent outer shell
layer.

Protoconch 350 ixm wide, finely granulate.

Teleoconch of up to 7.0 whorls, shoulder
and periphery strongly angulated by prominent
keels, ramp shallowly concave, shallowly concave
between periphery and outermost basal spiral,
more deeply concave between shoulder keel and
periphery ; base gently contracted, convex. Shoul¬
der keel summit narrowly rounded, strongly
serrate, becoming slightly higher than weakly
serrate peripheral keel al maturity, at about
adapical third ; subsutural angulation commencing
on 2nd whorl, gradually enlarging, becoming
serrate. Very minutely granulate throughout.
Axial riblets sigmoidal, narrow, strongly raised,
relatively very strong on spire, weaker on base,
weakening towards umbilicus, coalescing to form
strong spiral thread on adapical side of shoulder
keel, numbering 6 per mm between keels at end
of 5th whorl. Spiral threads crisp, considerably
finer than axials, multiplying by intercalation,
covering spire and base to outermost basal
spiral, commencing between outermost 2 basal
spirals on last adult whorl. Base with 9 or
10 crisp spiral cords, outer 3 or 4 spirals angulate
in section, inwardly decreasing in prominence,
innermost spiral weakest, bordering umbilicus.
interspaces considerably wider than each spiral.
Umbilical diameter 11.3 % of adult shell diame-
ter. Outer lip broken back, from growth Unes
posterior notch probably very deep ; basal notch
concave ; peripheral notch shallow, concave,

angulate. Pariétal glaze thin. Inner lip rather
thick, rim tightly folded towards umbilicus,
weakly flexed at base to form small, blunt, solid
tooth.

Animal unknown.

Type data. — Holotype (5.00 x 3.55 mm,
7.0 tw) mnhn, and immature paratype mnhn :
Biocal, stn CP 72. Paratype (mnhn) : Biocal,
stn DS 59.

Distribution. Off southem New Caledo¬
nia, 2 100-2 650 m (dead).

Remarks. — Seguenzia levii is highly distinc¬
tive in its tall, narrowly conical spire, very
prominent axial riblets and strong, serrate shoul¬
der keel.

Etymology. — It is named after Claude Levi,
who was cruise leader of the 1985 Biocal
campaign.

Seguenzia engonia sp. nov.

Figs 253-257

Description. — Shell up to 3.25 mm high,
broader than high, thin, widely umbilicate, spire
0.76-1.06 x as high as aperture ; white. nacreous
through thin, translucent outer shell layer.

Protoconch 300-320 izm wide, finely granulate.

Teleoconch of up to 5.10 whorls, shoulder
keel at adapical third, peripheral keel very
strong. ramp and side shallowly concave, shal¬
lowly concave between periphery and outermost
basal spiral ; base sharply contracted, weakly
convex. A long though distinct, weakly serrate,
subsutural angulation commencing on 4th whorl.
Axial riblets fine, crisp. sigmoidal. traversing
spire, base and outer part of umbilical wall,
numbering 12 or 13 per mm between keels at end
of 5th whorl. Spiral threads crisp, similar to
axials, multiplying by intercalation, covering
spire and base to outermost spiral. Base with 7-
9 crisply defined spiral cords, outermost angulate
in section and most prominent, interspaces con¬
siderably wider than each spiral. Umbilicus deep.
rim angulate, diameter 28.6-34.1 % of adult shell
diameter. Aperture subrhomboidal. Outer lip
thin ; posterior notch deep, retraction depth 6.6-
7.4% and protraction depth 34.1% of shell
diameter ; slightly flared at apex ; basal notch U-

Source : MNHN, Paris

102

BRUCE A. MARSHALL

Figs 253-267. Genus Seguenzia : 253 - 257 , Seguenzia engonia, holotypc, 3.10 x 3.52 mm, 256 * 35, 257 x i()5. 258-262

S. stegastris, hololype, 2.70 x 2.80 mm, 261 x 45, 262 x ] 05. 263 - 267 , S. plu humides. holotypc, 2.73 x ^ ?g mm ’

266 x 45. 267 x 70.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

103

shaped at rim, scarcely retracted behind. Pariétal
glaze thin. Inner lip rather thick, rim tightly
folded towards umbilicus, deeply curved towards
umbilicus, strongly flexed at base to form strong,
rounded tooth, channelled below.

Animal unknown.

Type data. Holotype mnhn (3.10 x
3.52 mm. 5.1 tw) and 3 paratypes (2 mnhn,

1 nmnz) (3.25 x 3.60 mm, 5.1 tw ; 2.95 x
3.50 mm, 4.75 tw) : Biocal, stn DW 79.

Distribution. Off Ouvéa, Loyalty Islands,

1 320-1 380 m (dead).

Remarks. Seguenzia engonia is a very
distinctive species characterised by low spire,
sharply angulate periphery, broad umbilicus, and
strong tooth on the inner lip.

Etymology. — Angular (Greek).

Seguenzia platamodes sp. nov.

Figs 263-267

Description. Shell (holotype) 2.73 mm
high, slightly broader than high, thin, umbilicate,
spire 1.10 x as high as aperture; white, nacreous
through thin, translucent outer shell layer.

Protoconch 400 |xm wide, surface eroded.

Teleoconch of 4.1 whorls, shoulder and peri¬
phery angulated by blunt-edged keels, ramp
and side shallowly concave, shallowly concave
between periphery and outermost basal spiral ;
base rather suddenly contracted, weakly convex.
Shoulder keel at adapical third, a low serrated
subsutural angulation commencing on 4th whorl.
Very minutely granulate throughout. Axial riblets
fine, crisp, sigmoidal, traversing spire and base,
extending onto outer part of umbilical wall,
numbering 11 per mm between keels at end of
4th whorl. Spiral threads crisp, finer than axials,
multiplying by intercalation, covering spire and
outer base to outermost spiral. Base with 8 crisp,
similar spiral cords, interspaces considerably
wider than each spiral. Umbilicus deep, rim
angulate, diameter 24.4 % of shell diameter.
Aperture subquadrate. Outer lip thin, retraction
depth 4.1 % of shell diameter, protraclion depth
unknown (rim damaged) ; basal notch concave,
peripheral notch very slightly retracted. Pariétal

glaze thin. Inner lip thick, shallowly curved
towards umbilicus, not flexed at base, toothless.

Animal unknown (dried).

Type data. — Holotype mnhn (2.73 x
3.28 mm, 4.1 tw) : Biocal, stn DS 14.

Distribution. — Off Lifou, Loyalty Islands,

3 680-3 700 m (alive).

Remarks. — Seguenzia platamodes most clo-
sely resembles S. engonia sp. nov., from which it
differs in being larger relative to the number of
whorls. in having a much larger protoconch, in
the later appearance of the subsutural angula¬
tion, and in having more bluntly angulate sum-
mits on the shoulder and peripheral keels. Judging
from the simple apertural features and the large
protoconch, the holotype may be immature.
Accordingly, larger specimens may be expected
to develop a tooth at the base of the inner lip.

Etymology. — Flattened (Greek).

Seguenzia stegastris sp. nov.

Figs 258-262

Description. Shell (holotype) 2.70 mm
high, about as high as broad, thin, umbilicate,
spire 1.07 x as high as aperture ; white, nacreous
through thin, translucent outer shell layer.

Protoconch 310 (xm wide, minutely granulate.
granules coalescing to form single spiral thread
at summit.

Teleoconch of 5.25 whorls ; shoulder and
periphery angulated by sharp-edged keels of
similar size, ramp and side concave, concave
between periphery and outermost basal spiral :
base gently contracted, convex. Shoulder keel at
about adapical third, summit slightly upturned ;
a prominent, finely serrate subsutural angulation
commences on 3rd whorl. Axial riblets fine,
crisp, sigmoidal, collabral, traversing spire and
base to spiral thread within umbilicus. num¬
bering 13 per mm between keels on spire at
end of 5th whorl. Spiral threads crisp. similar.
numerous, multiplying by intercalation, covering
spire and base from periphery to outermost basal
spiral, commencing between outer 2 basal spirals
on last half of last adult whorl. Base with 6 crisp
spiral cords. outermost angulate in section, most

Source : MNHN, Paris

104

BRUCE

MARSHALL

Figs 268-275. - Radulae and jaws (275) : 268-270, Asthelys nitidula. NE of Sandy Cape. Queensland (ams c. 154369),
268 x 830, 269 x 1 000, 270 x 1 250. 271, Ancistrobasis scilula. paratype. Bloc ai. stn DW 51. x 1 060. 212-215,

Ancistrobasis bouche!i. paratype. Biocal stn DW 77, 272 x 950. 273 x | 380, 274 x 1 210. 275 (jaws) x 150.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

105

Figs 276-283. Jaws (276, 277, 279) and radulac : 276, 277, Ancistrobasis boucheli, paratypc. Biocal stn DW 77 (jaw details).
276 » 700, 277 * 2 830. 278, 279, Calliobasis spectrum, paratypc. Biocal stn DW 41. 278 x 1890. 279

(jaws) x 220. 280, 281, Fluxinella asceta, paratypc. Biocal stn DW 33. 280 x 1 140. 281 x | 140. 282, 283.

Halystina caledonica, paratypc. Biocal stn DS 04, 282 x i 420, 283 x i 960.

Source : MNHN, Paris

106

BRUCE A. MARSHALL

prominent, widely separated from next spiral ;
inner spirals similar, doser though interspaces
considerably broader than each spiral, a fine
spiral thread in each interspace of 2nd-4th spirals
from umbilicus. Umbilicus deep, a spiral thread
on wall behind tooth on inner lip, diameter
22.6 % of shell diameter. Aperture subrhomboi-
dal. Outer lip thin ; retraction depth of posterior
notch 12.5% of shell diameter, protraction
depth unknown (labial projection broken) ; basal
notch concave ; peripheral notch small, angulate.
Pariétal glaze thin. Inner lip rather thick, rim
tightly folded towards umbilicus, deeply curved
towards umbilicus, sharply flexed at base to
form strong angulate denticle, channeled below.

Animal unknown (dried).

Type data. Holotype mnhn (2.70 x
2.80 mm, 5.25 tw) : Biocal, stn DW 80.

Distribution. Off Ouvéa, Loyalty Islands
900-980 m (dead).

Remarks. Seguenzia stegastris most closely
resembles S. eutyches sp. nov. in shape, and
differs primarily in having much stronger spiral
threads on the spire, and a considerably stronger
tooth on the inner lip.

Etymology. — A weaver (Greek).

Figs 284-287. Radula, Basitissa superba. Biocal stn CP 13, 284 x 508, 285 x l 164, 286 x i 154, 287 (showing
intcrlocking marginal tooth bases) x j 164. 6

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

107

DISCUSSION

The seguenziid fauna off New Caledonia and
the Loyalty Islands is exceptionally diverse in
comparison with the 89 Recent taxa hitherto
known from the rest of the world (Quinn, 1983b,
Table 1 ; Table 25 herein). Of the 55 species
présent, 50 (91 %) are unknown from elsewhere,
28 (51 %) are known from single stations, while
12 (22 %) are represented by single specimens,
the two latter totals suggesting that additional
species remain to be discovered. Judging from
Personal examination of rich unworked collec¬
tions from off the Philippine Islands (mnhn,
usnm), seguenziid species richness in this area
will probably prove to be at least as high.

Tablh 25. Géographie distributions of Recenl Seguen-

ziidac with numbers of nominale taxa. Numbcrs of species

shared with other régions in parenthesis.

New Caledonia and Loyalty Islands - 55 (6)

New Zealand région - 22 (4)

Western Atlantic 18 (2)

Indonesia, Malaysia, Coral Sea 12 (2)

Eastern Atlantic — 10 (1)

Northeaslem Pacific 7
Japan 6

Australia — 5 (2)

Soulhwestern Indian Océan and South Africa 5
Southern Océan and Antarctica — 3 (1)

Northern Indian Océan 2
Central Atlantic — 1
Northern Pacific I

The relatively low seguenziid diversities repor-
ted from elsewhere in the world Australia (5
species) and Japan (6) are particularly anoma-
lous may reflect insufficient sampling with

appropriate gear (i.e. fine-meshed epibenthic
sledges) at bathyal and abyssal depths. The
family is very poorly represented both as taxa
and individuals in extensive (ams) collections of
mollusca from depths shallower than 1 000 m otf
Australia, however, and species richness in this
depth range may not in fact be as high. By
comparison, 15 species from off New Caledonia
and the Loyalty Islands hâve mean depth occur¬
rences of less than 1 000 m (overall mean 709 m),
and four of these taxa are locally abundant.

Biocal samples from deep water (> 200 m)
o(T New Caledonia and the Loyalty Islands

contain exceptionally rich (largely unworked)
mollusc faunas in general, and to judge from the
remarkable Pleistocene assemblage at Santo,
Vanuatu (Ladd, 1976 ; 1982), il seems likely that
similarly diverse faunas will be found to occur
throughout Melanesia. Since the Melanesian arc
is situated at current or former boundaries of the
Australian and Pacific lithospheric plates, species
richness there is probably due at least partly to
progressive accumulation of taxa transported on
the plates.

Six taxa recorded from off New Caledonia and
the Loyalty Islands occur elsewhere in the western
Pacific : Basilissa superba (northern Coral Sea
and Philippine Sea), Asthelys nilidula (Queens¬
land), Ancistrobasis monodon (Malaysia), Quin-
nia panda (New Zealand), Seguenzia chelina
(New Zealand), and S. malara (New Zealand).
Basilissa superba (2 560-3 740 m) probably has a
continuous distribution through the Coral Sea to
New Caledonia, but is bathymetrically isolated
from the Philippine Sea population by island
arcs and associated trenches, which probably
accounts for the différence in umbilical morpho-
logy between the two populations. Known popu¬
lations of Asthelys nilidula (1 320-1 620 m), Ancis¬
trobasis monodon (505-680 m), Seguenzia chelina
(807-1 029 m), and S. matara (750-1 029 m) are
currently separated by depths considerably greater
than the deepest known occurrences of living
specimens. Asthelys nitidula and Ancistrobasis
monodon may hâve essentially continuous distri¬
butions via drifting eggs or larvae from popula¬
tions off the intervening islands, reefs and sub¬
marine banks, either through the Melanesian
Arc, or between Queensland and New Caledo¬
nia. That their distributions may be relicts of
formely more continuous distributions, rather
than the resuit of continuous larval dispersai
events, is suggested by différences in shell and
umbilicus size between the widely separated
populations. Known populations of Quinnia
patula (1 760-2 740 m), Seguenzia chelina and S.
matara are separated by the 1 400 km long
Norfolk Ridge, which is virtually continuous
between New Caledonia and New Zealand on
the 1 500 m contour, with a chain of rises on the
1 000 m contour (“ Gebco ", 1982). Although ail

Source : MNHN, Paris

108

BRUCE A. MARSHALL

of these species could easily hâve continuous
gene flow along the Norfolk Ridge, northern and
Southern populations of S. chelina and S. matara
may now be isolated by bathymetry.

Available evidence indicates thaï New Caledo-
nia and New Zealand had very similar geological
historiés throughout the Mesozoic and into the
Paleocene and Eocene (Grant-Mackie, 1985;
Lillie & Brothers, 1970 ; Stevens. 1980). Total
or partial emergence of the Norfolk Ridge
during the Mesozoic and perhaps into the Ter-
tiary could account for some of the observed
similarities in the présent day terrestrial biotas of
New Caledonia and New Zealand (Dawson,
1963; Stevens, 1980).

Evidence of major Eocene subsidence of the
northern part of Norfolk Ridge has been discussed
by Daniel et al. (1976) and Bitoun & Recy
( 1982), who respectively deduced subsidences of
about 400 m during the late Miocene or Pliocène

and about 1 500 m during the Miocene. If the
Southern part of the Norfolk Ridge subsided
during the same timeframe, this could account
for the disjunct distributions of S. chelina and
S. matara.

Potential biogeographical affinities of the New
Caledonian Recent marine fauna should be
sought in the subtropical mid tertiary faunas of
northern New Zealand, such as in the richly
fossiliferous lower Miocene beds at Pakaurangi
Point, Kaipara Harpara Harbour, and especially
Parengarenga Harbour. From a general biogeog¬
raphical standpoint it is appropriate to indicate
that the différences between the marine and
terrestrial biotas of New Caledonia and New
Zealand might reasonably be expected to be
almost as great as they are today if there was
currently continuous land between them, given
the latitudinal range covered.

ACKNOWLEDGEMENTS

I am grateful to P. Bouchet (mnhn) for the
opportunity to study this remarkable fauna. For
the loan of type material and reference specimens
I thank I. Hayami (Geological Institute, Uni-
versity of Tokyo), R. S. Houbrick (usnm),
A. Matsukuma (nsmt), R. G. Moolenbeek
(zma), A. Warên (Swedish Muséum of Natural
History, Stockholm), and K. M. Way (bmnh).
For discussions on south-west Pacific geology
1 thank H. J. Campbell, R. A. Cooper and

G. R. Stevens (New Zealand Geological Survey,
Lower Huit). Thanks to J. W. Dawson (Victoria
University, Wellington) for information on New
Caledonian plant biogeography, K. Downie and
D. Wakelin (nmnz) respectively for photo¬
graphie printing and word processing, and to
B. Burt and W. St George (New Zealand
Geological Survey, Lower Huit) for assistance
with scanning électron microscopy.

REFERENCES

Barnard, K. H., 1963. — Contributions to the
knowledge of South African marine Mollusca. Part 4.
Gastropoda : Prosobranchiata : Rhipidoglossa, Doco-
glossa. Tectibranchiata. Polyplacophora. Soleno-
gastres. Scaphopoda. Ann. S. Afr. Mus., 47 : 201-
360.

Barskov, I. S., Golovinova. M. A. & Goryachev, V.
N ., 1980. Structure of the nacreous layer of deep-

water molluscs of the genus Seguenzia (Mollusca,
Gastropoda). Dok. Akad. Nauk. SSSR, 252 : 1015-
1017.

Bayer, F. M., 1971. — New and unusual mollusks
collected by r/v John Elliot Pillsbury and r/v Gerda
in the tropical western Atlantic. Bull. Mar. Sci., 21 :
111-236.

Bitoun, G. & Recy, J.. 1982. Origine et évolution
du Bassin des Loyauté et de ses bordures après la
mise en place de la série ophiolitique de Nouvelle-
Calédonie. Trav. Doc. O.R.S.T.O.M.. 147 : 505-539.

Cernohorsky, W. O., 1978. - Tropical Pacific
Marine Shells. Pacific Publications, Sydney. 352 pp.

Coan, E., 1964. A proposed révision of the
rissoacean families Rissoidae. Rissoinidae, and Cin-
gulopsidae (Mollusca : Gastropoda). Veliger, 6 ■
164-171.

Dall, W. H., 1889. Reports on the results of
dredging under the supervision of Alexander Agas-
siz. in the Gulf of Mexico (1877-78) and in the
Caribbean Sea (1879-80), by the U.S. Coast Survey
Steamer "Blake" Lieut. - Commander C. D. Sigsbee,
U.S.N., and Commander J. R. Bartlett, U.S.N.]
commanding. 29. Report on the Mollusca. Part 2]
Gastropoda and Scaphopoda. Bu/l. Mus. Camp
Zoo!., 18 : 1-492.

Daniel, J., Dugas, F.. Dupont, J., Jouannic, C.,
Launay, J., Monzier, M. & Recy, J.. 1976. La
zone charnière Nouvelle Calédonie-Ride de Norfolk

Source : MNHN, Paris

MOLLUSCA GASTROPODA : SEGUENZIIDAE

109

(S. W. Pacifique) — Résultats de dragages et
interprétation. Cah. O.R.S.T.O.M.. Geol., 8 : 95-
105.

Dautzenberg, P. & Fischer, H., 1897. Campagnes
scientifiques de S.A.S. le Prince Albert 1" de
Monaco. Dragages effectués par 1‘Hirondelle et par
la Princesse-Alice. Mèm. Soc. Zool. France. 10 : 139-
234.

Dawson. J. W.. 1963. - New Caledonia and New
Zealand — A botanical comparison. Tualara. 11 :
178-193.

“Gebco", 1982. - Chart 5.10 (General Bathymétrie
Chart of the Océans). Canadian Hydrographie
Service, Ottawa.

Goryachev, V. N„ 1987. — The volume and the
position of the Family Seguenziidae (Mollusca,
Gastropoda, Seguenziidae) in the gastropod class.
In : Y. I. Starobogatov, A. N. Golilov, and
I. M. Likharev (eds), Molluscs, results and perspec¬
tives of investigation. Eighth meeting on the investi¬
gation of molluscs, USSR Academy of Sciences
Zoological Institute : 21-23.

Grant-Mackie, J. A., 1985. — New Zealand - New
Caledonian Permian - Jurassic faunas, biogeogra-
phy and terranes. N.Z. Geol. Surv., 9 : 50-52.

Iredale, T., 1917. On some new speeies of marine
Mollusca from Christmas lsland, Indian Océan.
Proc. Malac. Soc. Lond.. 12 : 331-334.

Jeffreys, J. G., 1876. Preliminary report of the
biological results of a cruise in H. M. S. Valorous to
Davis Strait in 1875. Proc. Roy. Soc.. 25 : 177-230.

Ladd, H. S., 1976. New Pleistocene Neogastropoda
from the New Hébrides. Naulilus. 90 : 127-138.

Ladd, H. S., 1982. — Cenozoic fossil mollusks from
Western Pacific isiands ; gastropods (Eulimidae and
Volutidae through Terebridae). U. S. Geol. Surv.
Prof. Pap.. 1171 : 1-100.

Lillie, A. R. & Brothers, R. N„ 1970. The
geology of New Caledonia. New Zealand J. Geol.
Geophys.. 13 : 145-183.

Marshall. B. A., 1979. The Trochidae and

Turbinidae of the Kermadec Ridge (Mollusca :
Gastropoda). New Zealand. J. Zool.. 6 : 521-552.

Marshall. B. A., 1983. Recent and Tertiary

Seguenziidae (Mollusca : Gastropoda) from the
New Zealand région. New Zealand. J. Zool.. 10 :
235-260.

Marshall, B. A., 1988. New Seguenziidae (Mol¬
lusca : Gastropoda) from the Tasman, South Pacific
and Southern Antilles Basins. New Zealand. J.
Zool.. 15 : 235-247.

Okutani, T.. 1964. Report on the archibenthal and
abyssal gastropod Mollusca mainly collected from
Sagami Bay and adjacent waters by the R. V. Soyo-
Muru during the years 1955-1963. J. Fac. Sci. Univ.
Tokyo. (2), 15 : 371-447.

Okutani, T., 1982. Rediscoveries of an abyssal
trochid, Basilissa superba Watson from the south of
Japan. Venus. 40 : 237-239.

Ponder W. F., 1967. — The classification of the
Rissoidae and Orbitestellidae with descriptions of
some new taxa. Trans. Rov. Soc. N. Z., Zool., 9 :
193-224.

Ponder W. F., 1985. — A review of the généra of the
Rissoidae (Mollusca : Mesogastropoda : Rissoacea).
Rec. Ausi. Mus., Suppl. 4, 221 pp.

Quinn, J. F., 1983a. — Carenzia. a new genus of
Seguenziacea (Gastropoda : Prosobranchia) with
the description of a new speeies. Proc. Biol. Soc.
Wash.. 96 : 355-364.

Quinn, J. F., 1983b. A révision of the Seguenziacea
Verrill, 1884 (Gastropoda : Prosobranchia). 1. Sum-
mary and évaluation of the superfamily. Proc. Biol.
Soc. Wash.. 96 : 725-757.

Quinn. J. F.. 1987. A révision of the Seguenziacea
Verrill. 1884 (Gastropoda : Prosobranchia). 2. The
new généra Hadroconus. Rolellenzia. and Asthelvs.
Naulilus, 101 : 59-68.

Salvini-Plawen, L. & Haszprunar. G.. 1987. The
Vetigastropoda and the systematics of streptoneu-
rous Gastropoda (Mollusca). J. Zool. Lond.. 211 :
747-770.

Schepman. M.. 1908. — The Prosobranchia of the
Siboga Expédition. Part 1 : Rhipidoglossa and
Docoglossa. Siboga-Expeditie. 49a : 1-107.

Schepman, M., 1909. The Prosobranchia of the
Siboga Expédition. Part 2 : Taenioglossa and Pteno-
glossa. Siboga-Expeditie. 49b : 109-231.

Stevens, G. R.. 1980. New Zealand Adrift. Read,
Wellington. 442 pp.

Thiele, J., 1925. — Gastropoda der Deutschen Tiefsee-
Expedition 2. Deutsche Tiefsee-Expédition. 1898-
1899. 17 (2) : 35-382.

Thiele, J., 1929-1935. — Handbuch der systematis-
chen weichtierkunde. 2 vols. Fischer. Jena. 1134 pp.

Verrill, A. E., 1884. — Second catalogue of Mol¬
lusca reccntly added to the fauna of the New
England coast and the adjacent parts of the Atlan¬
tic, consisting mostly of deep-sea speeies. with notes
on others previously recorded. Trans. Connect.
Acad. Sci.. 6 : 139-294.

Watson, R. B.. 1879. Mollusca of H.M.S. ‘Chal¬
lenger ' Expédition. 3. Trochidae, viz. the généra
Seguenzia, Basilissa. Gaza and Bemhix. J. Linn. Soc.
Lond. Zool., 14 : 586-605.

Watson, R. B.. 1886. Report on the Scaphopoda
and Gasteropoda collected by H.M.S. Challenger
during the years 1873-76. Repts Sci. Res. Challenger
Exp.. Zool., 42 : 1-756.

Wenz, W., 1938-44. — Gastropoda. Allgcmeinc Teil
und Prosobranchia. In : O. Schindewolf (ed.),
Handbuch der Palaozoologie, 6 (1). 2 vols. 1639 pp.

Source : MNHN, Paris

Source : MNHN, Paris

ULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSUL'

4

Mollusca Gastropoda : Systematic position and révision
of Haloceras Dali, 1889
(Caenogastropoda, Haloceratidae fam. nov.)

Anders WARÉN

Naturhistoriska Riksmuseel
Box 50007
10405 Stockholm
Sweden

Philippe BOUCHET

Muséum national d'Histoire naturelle
Laboratoire de Biologie des Invertébrés marins et Malacologie
55, rue Buffon
75005 Paris
France

ABSTRACT

Haloceras. until now included in thc Trichotropidae, is
made the type of a ncw Family, Haloceratidae. considercd
related to but more primitive lhan the Tonnoidea and the
Echinospirida, on the basis of new anatomical information.
The main différence from existing tonnoidean families is a
more primitive nervous System and the absence of complex
salivary glands. The new family shows similarities to the
Trichotropidae. a family which should be included in the
Echinospirida. Micropiliscus Dali. 1927, until now included in
thc Trochidae, is synonymized with Haloceras. and the new

genus Zygoceras is introduced. Seventeen named species (ten
new) and thrcc left unnamed are here included in the family.
Ail species live on the outer part of the continental shelf and
the slope. down to about 3500 m depth. Species of Haloce-
ratidac are known from the mid and low latitudes of the
Atlantic, lndian, East and West Pacific océans. Ail but one
species hâve planklotrophic larval development and one
species is known from both the Atlantic and thc lndian

RESUME

Position systématique et révision du genre Haloceras Dali,
1889 (Caenogastropoda, Haloceratidae fant. nov.).

Le genre Haloceras était jusqu'ici classé dans les Tricho-
tropidac. Sur la base de nouveaux caractères anatomiques,u-
ne famille nouvelle Haloceratidae est créée. Elle est considé¬
rée proche de, mais plus primitive que les Tonnoidea et les
Echinospirida. Les différences principales qui séparent les
Haloceratidae de la lignée tonnoïde sont un système nerveux
plus primitif et l'absence de glandes salivaires complexes.
Cette nouvelle famille présente des ressemblances avec les
Trichotropidae, qui doivent d'ailleurs être classés dans les

Echinospirida. Le genre Micropiliscus Dali, 1927. jusqu'ici
classé dans les Trochidae. est placé en synonymie d‘ Haloce¬
ras. et le nouveau genre Zygoceras est décrit. La famille
comprend dix-sept espèces nommées (dont 10 nouvelles) et
trois laissées sans nom; toutes vivent sur les parties profondes
du plateau continental et sur la pente, jusqu'à 3500 m
environ, aux latitudes basses et moyennes dans les océans
Atlantique. Indien, Ouest et Est Pacifique. Toutes les espèces,
sauf une. ont un développement larvaire planctotrophe et une
espèce est connue à la fois de l'océan Atlantique et de l'océan
Indien.

Warbn. A. & Bouchet, P.. 1991 Mollusca Gastropoda : Systematic position and révision of Haloceras. Dali 1889 (Caenogastropoda.
Haloceratidae fam. nov.). In : A. Crosnii-r & P. BoucmrT (eds). Résultats des Campagnes Musorstom, Volume 7. Mèm. Mus. nam. Hisi. nui..
(A), 150 : Il 1-161. Paris ISBN : 2-85653-180-6.

Publié le 20 mars 1991.

Source : MNHN, Paris

112

ANDF-RS WARÉN & PHILIPPE BOUCHET

INTRODUCTION
Historjcal background

Based on shell and opercular features, Dali.
(1889) introduced the name Haloceras for lhe
NW Atlantic deep-sea gastropod Cithna cingu-
lata Verrill, 1884. He considered Haloceras a
subgenus of Separatista Gray, 1847 (then placed
in the Adeorbidae, now, in our view correctly,
placed in lhe Trichotropidae). Thiele (1929: 243)
and Wenz (1940: 890) classified Haloceras as a
subgenus of Torellia Jeffreys, 1867 in the family
Trichotropidae ( = Capulidae).

Dall (1927) described Micropiliscus, as a
subgenus of the trochid genus Solariella Wood,
1842, differing from the typical species by having
a brownish larval shell. Micropiliscus has since
been retained in the Archaeogastropoda as a
subgenus of Solariella by the few authors that
hâve menlioned or listed il (Thiele, 1929: 48;
Wenz, 1938: 275; Quinn, 1979: 43).

During the last 10 years we hâve been aware
that there exist several species similar to Haloce¬
ras, and we hâve been accumulating material of
them and searching muséum collections we hâve

The genus Cithna A. Adams, 1863 has fre-
quently been used for strange. globular or dis-
coid gastropods, among them Haloceras. Pre-
sently (Thiele, 1929: 64; Wenz, 1938: 336)
Cithna is classified among the archaeogastro-
pods. We figure (Figs 44-45) a syntype from
USNM, to show the planktotrophic larval shell

Abbreviations used in text:

sh(s) - - (empty) shell(s);
spm(s) — (live taken) specimen(s);
ams The Australian Muséum, Department of
Invertebrate Zoology, Sydney:
bmnh British Muséum of Nalural History,
Mollusca Section, London;
lacm Los Angeles County Muséum of Natu¬
rel History, Los Angeles;
mnhn Muséum national d'Histoire naturelle.
Paris;

visited. The présent paper is based on ail the 103
specimens and shells that, to our knowledge,
hâve been taken by océanographie expéditions
worldwide in 120 years. The absence of fossil
record should probably be inlerpreted as a
reflection of the rarity of the family. It is
remarkable to note that. whereas the first halo-
ceratid specimen was collected in 1870 (although
described in 1883), 64 % of lhe specimens now
available hâve been taken in just the last 20
years. No haloceratid was taken during the
“ Challenger ", “ Valdivia " or " Siboga " expé¬
ditions. and half of the species had not been
collected 10 years ago. Two régions, the South¬
west Pacific with 8 species and the North
Atlantic with 6 species. now contain together
82% of the species of the family, but this simply
represents a collecting effort artefact. We believe
thaï future exploration of tropical deep waters
will greatly increase the number of species in lhe
family.

A. Adams, 1863

which directly excludes Cithna from the Ar¬
chaeogastropoda. The soft parts are not known,
but a position in lhe vieinity of Scrupus Finlay,
1924 (position in Rissooidea presently unknown,
see Ponder, 1985), seems very likely. judging
from shell morphology.

mom Musée océanographique, Monaco;

nmnz National Muséum of New Zealand,
Wellington;

nsmt National Science Muséum, Tokyo;
rmnii Rijksmuseum van Natuurlijke Historié,
Leiden;

usfc United States Fisheries Commission;
usnm U.S. National Muséum of Naturel
History, Division of Mollusks, Washington
D.C.:

zmc Zoologisk Muséum. Kôbenhavn.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOŒRAT1DAE

113

MATERIAL AND METHODS

Most species are known from empty shells,
sometimes with partly decayed and dried soft
parts. These were used for radular préparation
after rehydration. The alcohol prescrved spéci¬
mens turned oui to be a problem since the soft
parts could not be extracted without damaging
the shells. which was too great a drawback when
often only one or two shells were known of the
species. Therefore they were dried and the soft
parts loosened from the columella by gently
pushing them backwards, into the shell. After
thaï they were rehydrated in buffered formalin
and extracted. They could then be used for
examination of external morphology and some
rough dissections, including préparation of the
radula.

The anatomical descriptions are thus mainly

based on dried and rehydrated specimens of:
Haloceras carinata (rmnh)

Haloceras cingulata (usnm 52077)

Haloceras galerita (holotype). A partly damaged
head-foot was serially sectioned in the same
way as Z. tropidophora. It contained none of
the organs behind the cephalopedal haemo-
coel.

Haloceras japonica (lacm 67-167.2)

Haloceras tricarinata (" Discovery " stn 10141)
Zygoceras tropidophora (paratype). The specimen
was decalcified, serially sectioned at 5 pm and
stained with Ehrlich's haematoxylin - eosin.
One specimen was dried to allow safe removal
of the body from the shell after rehydration
and used for radular préparation.

ANATOMY

EXTERNAL MORPHOLOGY
Figs 1-2

The general colour of the soft parts in alcohol
is whitish-yellowish transparent, with a more
opaque osphradium and occasionally (Z. tropi¬
dophora) a brownish-violet intestine and sto-
mach.

The body consists of about 1.7 whorls in Z.
tropidophora , of which the voluminous head-foot
and palliai complex occupy 0.25-0.30 whorls.
The rather shallow palliai cavity reaches back
about 0.25 whorls and its most posterior part
forms a blunt wedge-shaped space between the
pericardium and cephalo-pedal haemocoel.

The columellar muscle (Fig. 2) is solid and
broad but very short.

The head-foot is large, occupies about 0.25
whorls (slightly more in the tall-spired species)
and the palliai skirt covers most of this when the
soft parts are retracted. Many of the internai
organs can be seen by transparency (Figs 1-2). In
H. tricarinata the posterior part of the foot is
quite suckershaped. in other species less so but to
some extent it is divided in a more solid.
rounded, posterior part and an anterior, thinner,
strongly folded, and probably very motile part.
The latter has been strongly contorted in ail

specimens but its anterior dorsal side carries a
large and fleshy propodium, well aligned with the
anterior edge of the foot. The lower part of the
side of the foot is distinctly demarcated from the
upper parts, by a more or less distinct horizontal
furrow (Fig. 3) and by having a rougher surface.
H. tricarinata and Z. tropidophora hâve a large
posterior pedal gland opening in the posterior
part of the foot. This could not be discerned in
the other species, but neither could its absence be
ascertained. On the rear, dorsal part of the foot,
there is a thick but fragile, yellowish operculum.
In species with a depressed or disc-shape shell the
operculum (Figs 28, 31-34) is fan-shaped with the
initial coils situated apically, but they are usually
worn off in large specimens, so only the lalest
formed section of the fan remains. In more
tall-spired species the operculum is more ovate
and spirally coiled. If the nucléus is left, there is
a distinctly set off part, which is thinner. smoo-
ther and multispiral, and which corresponds to
the larval operculum.

The head is about as broad as the foot and
equipped with tentacles which in one specimen of
Zygoceras tropidophora are tapering, long and

Source : MNHN, Paris

114

ANDERS WAREN & PHILIPPE BOUCHET

Figs. 1-2. Zvgoceras tropidophora, extcrnal view ol soit
paris. I. 'right side. 2. Lefi side. Fine vertical linos
indieate position of cross sections with corresponding
Fig. number.

aa antenor aorta; au auriclc; bv blood
vessel; cm columcllar muscle: e — eye: i
intestine; k rénal organ; me mantle edge; mp
mesopodium; ng rénal gland; o opcreulum; oe -
ocsophagus; p pénis; pu posterior aorta; ph
proboscis; pp - propodium: r rectum; si
stomach: te tcntacle; ve vcntriclc.

slender; in the other species, strongly wrinkled,
short and conical, and evidently strongly
contracted. AU species hâve large eyes situated
dorso-laterally on, and of a diameter correspon¬
ding to about 2/5 of. the tentacle base. The
tentacles are connected by a thin skin-lold.
covering the proboscis. The dorsal body-wall is
quite thick and muscular.

Presumably no species hâve a snout. This
could, however, not be veriiied in some spéci¬

mens, because the proboscis has been more or
less everted in ail specimens examined and in
some specimens which were badly preserved and
dried it could not be seen if there was a tip of a
proboscis or a snout.

The pleurembolic proboscis, often partly ever¬
ted, is quite solid and muscular, in Z. tropido-
phorei 5 mm long and 2.5 mm diameter, but can
unquestionably be extended much more since the
wall of the proboscis sheath is about 1 mm thick.
The buccal mass is fairly small, 0.8 mm long and
0.6 mm diameter. There is a pair of solid. slender
jaws (Figs 4. j; 29, 30).

AU specimens examined had a simple, flatte-
ned, finger-shaped pénis altached slightly behind
and below the right tentacle (Fig. 1). In H.
japonica, II. galerita and H. cingulata, the pénis
also had an anteriorly dirccted simple fold-like
process. Such a flap was missing in Z. tropido¬
phora and H. carinata. It was not possible to
distinguish an external sperm groove, but il can
be seen in sections of the two sectioned species.

Some species (H. cingulata. Z. tropidophora)
hâve an external skinfold on the inner and distal
part of the palliai oviduct, but ils morphology
could not be studied in detail.

One specimen of Haloceras carinata had a
large egg-mass in the palliai cavity (Figs 64-65),
measuring 0.9 x 0.9 x 1.6 mm and consisting of
about 200 developing, shell-less embryos of a
diameter of about 0.16 mm. Due to the dried
State of the specimen it was not even possible to
verify the presence of an oviduct. but a pénis was
présent.

One specimen. Haloceras japonica. proved to
be ovoviviparous and had 9 large embryos (Fig.
64) in the partly ruptured (from drying?) ovi-
ducl. These embryos were lying in the duct
behind the palliai cavity. along almost one whorl
of the viscéral mass. This specimen also had a
pénis of normal appearance and position for the
genus.

The palliai cavity is wide and spacious (Fig. 3).
The anterior edge of the palliai skirt has a simple,
thick and muscular. well delined zone with no
papillae or appendices. Al the left corner this
zone is broader and thicker. and may function as
a siphon.

The gill occupies a narrow zone, from the
thickened edge of the palliai skirt backwards to
the bottom of the cavity and consists of about 25
flattened. lingershaped Icaflets, in Z. tropido-

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NF.W FAMILY HALOCERAT1DAK

115

Fig. 3. Zygoceras tropidophora, cross section of body behind tcntacles.

apg acccssory pcdal ganglion: av alïereni branchial vcssel (Iront rectal sinus); bmr buccal mass retractor:
epe ccrebro-pcdal connective; a etenidium; dpg duel of posterior pcdal gland; hg hypobranchial gland:/g
latéral groove; mb muscle bundlcs l'rom proboscis sheath: oc ocsophagus: og osphradial ganglion: os
osphradium; p pénis: pm palliai margin; pn - (outer) pcdal nerve: poil palliai oviduct; ppg posterior pcdal

gland; reg right cérébral ganglion; rm - - retractor muscle of proboscis: rpg righl pcdal ganglion: rs rectal sinus;

sia — statocyst. Scalc line I mm.

Source : MNHN, Paris

116

ANDERS WARÉN & PHILIPPE BOUCHET

phora 1.1 mm long and of a diameter of aboul
0.1 mm. In H. galerita they were 45 in number.
0.5 mm long. 0.07 mm broad and 0.03 mm thick.
In the posterior part of the cavity the gill is fairly
central, anteriorly it bends slightly over to the
left.

The osphradium is partly bipectinate and it is
situated along the anterior 3/5 of the gill. Il
consisls of 15-30 leaflets (in Zygoceras tropido-
phora 0.5 mm long and 0.12 mm broad) facing
the gill and half that number of smaller leaflets
along facing to the left. In H. tricarinata only the
outer half was bipectinate, in Z. topidophora the
inner half bipectinate and in H. cingulata il was
probably monopectinate.

The rectum runs almost centrally in the palliai
cavity and ends close to the edge of the palliai
skirl. Ils most distal part is free from the palliai
roof.

The specimen of //. carinata had white faecal
pellets in rectum and a foraminiferan in the
slomach. A specimen of Z. tropidophora had a
few small faecal pellets in rectum, consisting of
unidentified organic material and free from mi¬
nerai parlicles.

The hypobranchial gland is very thin and
inconspicuous. It covers the inner side of the
rectal sinus and a part of the palliai skirt in the
central part of the palliai cavity. but can only be
discerned in histological sections.

INTERNAI. ANATOMY
Figs 3-27

Head-foot

The tentacles, the sides and the back of the
head-foot are covered by a thin epithelium, on
the sides of the foot with numerous large mucus
producing cells mixed with the épithélial cells.
Mucous cells are common also on the pénis. The
hypobranchial (Figs 3, 5, hg) gland consists of a
single layer of sccreting cells, which covers the
left side of the rectal sinus and the palliai roof
leftwards to the gill, from just in front of the
rectum back to the kidney.

The sole of the foot has a dense layer, one cell
thick, of mucous cells directly under the epithe¬
lium. The posterior pedal gland (Fig. 3, ppg) is
large and voluminous, anteriorly bilobed and
abutting the pedal ganglia. Less than half-way to
its opening the two lobes unité and form a
common duct (Fig. 3, dpg) which opens poste-
riorly and centrally via a wide pore. The anterior
pedal gland (Fig. 6, ang) fills a large cavity
between the pro- and mesopodium, invades them
both, and extcnds back to the pedal ganglia
where it abuts the posterior pedal gland. It has a
wide opening between the pro- and mesopodium.

A liment ary canal
Figs 4, 11-22

The alimentary System consists of a pleurem-
bolic proboscis. a buccal mass with taenioglos-

sate radula. an anterior oesophagus with latéral
pouches, a long midoesophagus, a comparatively
short posterior oesophagus, a simple stomach, a
short intestine and a long rectum.

The anterior part of the proboscis sheath is
solid and muscular and can be retracted so the
true mouth is situated immcdiately behind the
tentacles and the posterior end of the muscular
section abuts the cérébral ganglia. It is manipu-
laled by two major ventral muscle bundles (Fig.
3, rm) which connect il to the columellar muscle,
and numerous smaller muscles (Fig. 3, mb),
which in Z. tropidophora end in the walls of the
cephalopedal haemocoel. In H. galerita they fuse
with the two ventral muscle bundles. The rear
part of the sheath is extremely thin-walled (Fig.
5, pps) and when the proboscis is retracted, it
leads from the rearmost, thick-walled section
forwards to the level of the tentacles.

The mouth is situated terminally at the tip of
the proboscis (Fig. 4. mo). The sides of the oral
cavity are equipped with a pair of prismatic,
semicircular jaws (Figs 4. /: 29-30) and the buccal
cavity is largely lined with cuticle. The salivary
ducts open shortly behind the jaws dorso-
laterally via a small pore. Their ducts meander
backwards; in H. galerita ail the way through the
nerve-ring. in Z. tropidophora they stop halfway.
The salivary glands each consist of a single
thick-walled coiled tube (Fig. 4, sg).

In Z. tropidophora the radular sac exils poste-

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

Figs. 4-6. 4. Zygoceras tropidophora, horizontal section through anterior part of proboscis. 5. H. galerita, cross-section

Ihrough head-foot between tentaclcs and pénis. 6. H. tropidophora, horizontal section through pro- and
mesopodium.

ang — anterior pedal gland; apg acessory pcdal ganglion; /><• - buccal cavity; bev — buccal connective; hg
- buccal ganglia; dfc dorsal food channel: j — jaw; mo mouth; mp mesopodium; oc oesophagus; pp
propodium; ppg — posterior pedal gland; pps posterior part of proboscis sheath; pr proboscis; ps proboscis
sheath; rc radular cartilage; rsc radular sac; sg — salivary glands; so sole of mesopodium. Scale line 0.25 mm.

Source : MNHN, Paris

ANDERS WARÉN & PHILIPPE BOUCHET

Figs. 7-10. Hafoceras tropidophora, cross section of palliai cavity and viserai mass. Position of sections indicaled in Fie
2. 7. Middlc part of palliai cavity. 8. Anlerior part of kidney. 9. Most posterior part of palliai cavity. jn

Viscéral hump, anterior lo stomach.

au anterior aorta: abv afferent branchial vein; arv afferent rénal vein; au auricle; cf ctenidial
filament; cm columellar muscle; eph cephalo-pedal haemocoel; dp digestive gland; mg efferent cavity, collectine
blood from rénal gland and ctenidium; /' - intestine; is — intestinal sinus; k rénal organ; ko rénal opening- hr

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERAT1DAE

119

ieft viscéral connective: If latéral groove on free lobe of palliai oviduct; ng — rénal gland: <><• oesophagus; og
osphradial ganglion; on osphradial nerve; os — osphradium; pc — palliai cavity; peu pericardium; p - pénis;
poü palliai oviduct; r rectum; rs rectal sinus: rsc — receptaculum seminis: rvg right viscéral ganglion: seg

séminal groove; sk skclctal rod; .vv — séminal vesiclc; les — tcsticle: vc — viscéral connective: vc ventricle.

Scale Unes 7-8, 0.25 mm, 9-10. 0.5 mm.

Source : MNHN, Paris

120

ANDERS WARÉN & PHILIPPE BOUCHET

riorly and makes a sharp turn ventrally, then
dorsally, so it ends ventrally to the anterior
oesophagus, immediately behind the buccal
mass. In H. galerita it exits posteriorly and
parallels the anterior oesophagus (Figs 16-17,
rsc) for a short distance before it ends just in
front of the buccal ganglia, which are situated
unusually far back in this species. The radular
cartilages are well developed.

The oesophagus starts with a dorsal food
channel (Fig. 16, dfc), which as soon as the
ventral floor has been formed, widens to form a
pair of shallow and thin latéral pouches over the
posterior part of the buccal mass. These disap-
pear behind the buccal mass. In H. galerita the
dorsal food channel continues dorsally, ciliated
and glandular, while the ventral floor of the
oesophagus is lined by a cuticle (Fig. 17). The
dorsal folds become broader and are strongly
ciliated and the oesophagus gets a roughly
I-shaped cross section, ail the way back to the
nerve ring, where the oesophagus widens and the
walls become more uniform ail around. Here the
oesophagus is also quite compressed and folded
so it was not possible to discern the torsion. In Z.
tropidophora the ventral part of the anterior
oesophagus is not lined by a cuticle, but the
whole circumference, except the dorsal food
channel is lined by mucus producing epithelium
(Fig. 11). This thick-walled, ventral part then
splits in two parts connected by a thin wall. At
the same lime the dorsal food channel has
rotated to become ventral (Fig. 13, dfc) giving
rise to an I-shaped portion (Fig. 13).

The posterior oesophagus (Fig. 14) begins at
the end of the cephalopedal haemocoel and has
folded walls. It opens ventrally and slightly in
front of the posterior end of the stomach, jointly
with two openings from the digestive gland. It
could not be ascertained if these openings cor¬
respond to an anterior and posterior lobe of the
digestive gland.

The stomach (Fig. 2, st; 18) is situated close to
the left side of the body, just behind the pericar-
dium. It is simple and sausage-shaped. tapering
towards the anterior end where it goes over into
the intestine via the style-sac without any abrupt
change. It is almost three times as long as broad.
There seems to be no crystalline style.

To the right of and posteriorly to the oeso-
phageal opening there is a prominent gastric
shield and a large cuticularized area where food

is accumulated. In the ventral, posterior part of
the stomach run two Iow and broad typhlosoles
which continue some distance into the style sac
(Fig. 20, tl. t2).

The intestine leaves the anteriorly and dorsally
situated posterior part of the stomach, turns to
the right under the left-most and posterior
extension of the kidney (Fig. 10, /), in a very
spacious haemocoel (Fig. 10, is), then it enters
the palliai skirt and turns to the left, so the anus
is more centrally situated than the proximal parts
of the rectum (Fig. 7). There was no great
histological différence between the proximal and
more distal parts (rectum) (Figs 21-22).

Vascular System
Figs 7-10

Ail organs are lying in a System of very
spacious venous sinuses which are filled by a
spongc-like System of interconnected cells, giving
the tissues a very porous appearance.

The pericardium (Fig. 9, pca) is situated with
its anterior end above the posterior part of the
palliai cavity, with the auricle in front of and
above the ventricle. In cross-section it is roughly
triangular with one corner drawn out towards
the center of the whorl. The renopericardial duct
leaves this corner at the anterior part of the
ventricle. There is no gonopericardial duct.

We assume that blood from the cephalopedal
haemocoel collects in a System of large anasto-
mosing venous cavities (Figs 9, 23, arv) which
communicate with the viscéral and intestinal
haemocoels and the kidney. There is also a
somewhat separate dorsal vein (Fig. 23, A-v)
which starts from spaces in the intestinal haemo¬
coel and the venous cavities and connects with
the kidney folds and the rénal gland (Fig. 23, ng).

The efferent branchial vein and the efferent
rénal gland vein (Fig 23, eng) join anteriorly to
the pericardium and carry the blood to the
auricle.

The aorta leaves the ventricle posteriorly and
splits directly into one anterior and one posterior
branch. The posterior aorta is very thin-walled
and continues superficially backwards, sending
off smaller vessels which supply the viscéral
hump with blood. It was followed backwards
past the stomach. The anterior branch of the
aorta (Figs 8-10, ad) turns forwards, passes

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

121

Figs. 11-l l ). Anterior alimenlary canal and stomach of /.. tropidophora (11-14. 18. 19) and H. galerila (15-17). 11.

Anterior oesophagus with dorsal food channcl. 12-13. Midocsophagus. after the rolalion of dorsal food channcl.

14. Poslcrior oesophagus. 15. Proboscis lip with oral tube and lefl jaw. 16. Proboscis and buccal mass, rctractcd
into head-foot. 17. Anterior oesophagus between buccal ganglia and buccal niass. 18. Stomach. longitudinal
section al close to right angle to oesophagus and intestine. - 19. Detail of gastric shield.

bev buccal connective; br - buccal mass retractor; c — cuticle lining; dfc dorsal food channel; dg
digestive gland; dsg - duct of salivary gland; fo — food; gx — gastric shield; i position of exil of intestine from style
sac: / jaw; oc position of entrance of oesophagus; oev ocsophageal valve: rxc radular sac. Scale Unes 0.1 mm.

Source : MNHN, Paris

122

ANDERS WARÉN & PHILIPPE BOUCHET

Figs. 20-23. Zygoceras tropidophora, postcrior alimcntary canal and kidney. 20. Intestine shortly aftcr stomach 21
Intestine more distally. 22. Rectum. 23. Palliai skirl with antcrior part of kidney.

cf ctcnidia! filament: eng elTcrent cavity. collecting blood from rénal gland and gill. leading to auricle- A.

fond string; kf rénal folds; ko rénal pore; A r vessel connecting kidney folds and rectal sinus- ng , e j
gland: r rectum; r.v rectal sinus: sk skelelal rods in gill filament; il, i2 typhlosolcs; ing tubules or r.-n^i
gland. Scale lines 0.1 mm. nal

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

123

Figs. 24-26. 24. Z.

iropidophora, Distal,
closed part of palliai
oviduct. 25. Halo-
ceras galerila. cepha-
lic tentacle with eye.

26. Z. iropido¬
phora, opcn part of
palliai ociducl and
sperm grove.

ag — albumen
gland; cm - columcl-
lar muscle; If — laté¬
ral fold; ms palliai
skirt; mu -mucus pro-
ducing epithelium; pc
— palliai cavity; pg
pigment layer: pod
palliai oviduct; rd
rods of retina; le
cephalic tentacle.
Scale Unes 24. 26. 100
pm, 25. 50 pm.

through the anlerior left corner of the rénal
gland and enters lhe cephalopedal haemocoel to
the left of the oesophagus. It then follows the
oesophagus forwards, turns to the right to a
position dorsally to the oesophagus and splits
in several smaller vessels shortly behind the
supraoesophageal ganglion.

Excretory organs
Figs 2. 8-10, 23

The rénal organ extends back to just past the
ventricle: rightwards it extends dorsally and
ventrally to the intestine; anteriorly it invades the
palliai roof (Fig. 8, k). One lobe of the rénal

Source : MNHN, Paris

124

ANDF.RS WARÈN & PHILIPPE BOUCHET

gland protrudes in front of the pericardium,
surrounds the efferent branchial and rénal ves-
sels, and the anterior aorta and extends down to
the columellar muscle (Figs 8-10, ng).

The rénal pore (Fig. 23, ko) is situated dorsally
in the palliai roof just in front of the pericar¬
dium.

Dorsally the right half of the rénal organ is
occupied by rénal folds; the left half consists of
the rénal gland, penetraled by several narrow,
blindly ending ducts from the cavity of the
kidney (Fig. 23, tng).

Nervous System
Fig. 27

The nervous System is partly concentrated with
the pleural ganglia being fused with the cérébral
ones and short and thick commissures and
connectives in the nerve-ring. The oesophageal
ganglia hâve rather long, but solid, connectives
and the viscéral and buccal connectives are long
and very thin.

The pedal ganglia are about as long as high
and somewhat wider. They are connected by a
very short commissure. At their anterior and
ventral ends two large nerves emerge close to
each olher, an anterior one leading to an anterior
accessory pedal ganglion and a ventral one
leading to a posterior accessory ganglion. Both
these pairs of ganglia are partly embedded in the
posterior pedal gland. Each pedal ganglion also
sends out at two large nerves laterally. The
anterior accessory pedal ganglia innervate the
anterior parts of the foot. The posterior acces¬
sory ganglia each send out two nerve stems
backwards in the foot, of which the more central

Fie;, 27. Zygoceras tropidophora , nervous syslem. Pedal
ganglia separated from cérébral ones; only right
statocyst drawn.

aag — anterior accessory pedal ganglion; bev

- buccal connective; ca - commissure belween
posterior accesory pedal ganglia; di dialyncury; ipn

- inner pedal nerve; Ibg left buccal ganglion; leg
left cérébral ganglion; Ipg left pedal ganglion; Ipn
latéral pedal nerve; Ipn left palliai nerve; Ivg - left
viscéral ganglion; opn — outer pedal nerve; par-
commissure between posterior accessory pedal gan¬
glia; pag posterior accessory pedal ganglion; plg —
pleural ganglion; rpn right palliai nerve; rvg right
viscéral ganglion; shg — subeosophageal ganglion; spg

supraoesophageal ganglion; sic statocyst; in —
tentacle nerve; zy — zygoncury.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

125

nerve innervâtes the rear part of the foot, the
more latéral one the middle part and sides of the
foot. No connections between the stems were
found. There is also a large nerve from each
posterior accessory ganglion which follows the
duct of the posterior pedal gland backwards for
a short distance and then connect dorsally to the
duct.

The cérébral ganglia are lying abutting each
other, connected by a thin commissure. The right
ganglion is completely fused with the right
pleural ganglion. The tentacle nerve exits dorso-
laterally at the anterior 1/4 and the nerve is
simple. The buccal connective, together with
several other nerves to the proboscis, leaves
anteriorly. Posteriorly the right cerebro-pleural
ganglion is drawn out into the pleural-supra-
oesophageal connective and a palliai nerve for-
ming a zygoneury with the suboesophageal gan¬
glion. The left cérébral ganglion is separated
from the pleural ganglion by a shallow constric-
tion. The left pleural ganglion sends out the
pleural-suboesophageal connective and a palliai
nerve which forms a dialyneury with the osph-
radial nerve. The cerebro- and pleuro-pedal
connectives are of a length corresponding to 2/3
of the width of the combined cérébral ganglia
and the pleural connective is about twice as thick
as the cérébral one.

The suboesophageal ganglion is situated some
distance behind the nerve ring in the right corner
of the cephalopedal haemocoel. the supraoeso-
phageal ganglion slightly further back and more
dorsally. It sends out a large osphradial nerve.
which after the dialyneury continues in the palliai
skirt, anteriorly, till it joins the osphradial axis
which is occupied by the large osphradial gan¬
glion throughout its length.

The viscéral connectives are long and slender
and it was not possible to find the connection
between the two viscéral ganglia of which the left
one lies in the cephalopedal vein, the right one in
the space where the cephalopedal and viscéral
veins join.

In Z. tropidophora the buccal ganglia are
situated on the posterior part of the buccal mass
and joined by a commissure of about the same
length as the diameter of the ganglia. In H.
galerita the buccal ganglia are situated further
back. behind the radular sac. closely appressed to
each other and to the anterior oesophagus.

The innervation of the pénis could not be

worked out in detail, but it may be of both
cérébral and pedal origin.

Sensé organs

A pair of statocysts is présent, lying postero-
laterally to the pedal ganglia in Z. tropidophora
(Fig. 3, sta), posteriorly in H. galerita. They hâve
a single statolith.

The eyes (Fig. 25) are unusually large, equip-
ped with a lens and a well developed retina. They
are innervated by a branch of the tentacle nerve.

The osphradium (Figs 3, 7, os) is well develo¬
ped, bipectinate with more leaflets along the
inner side than on the outer side (see external
morphology). The axis is fdled by the large
osphradial ganglion (Fig. 7. og). Due to poor
préservation no detailed observations were made
on the structure of the leaflets except lhat they
are constricted basally and strongly ciliated
above the constriction.

Reproductive organs

Figs 3, 7-10. 24, 26

In the sectioned specimen of Z. tropidophora
the palliai oviduct is probably not fully formed,
but it possesses also a complété set of male
organs. These form the basis for the description.

The male organs consist of testis, a long and
winding sperm duct opening in the posterior part
of the oviduct; possibly a gutter along the free
lobe of the oviduct; and a pénis with a sperm
gutter.

The testis (Fig. 10, tes) is situated along the
right and more posteriorly also the dorsal side of
the viscéral mass. It consists of numerous tubules
which anteriorly and ventraily unité to form a
common sperm duct. which also functions as a
séminal vesicle. Beside eu- and paraspermatozoa.
there are also scattered eggs in early developmen-
tal stages in the gonad.

The sperm duct starts with a few coils (Figs
9-10, sv), then it passes straight forwards. Most
of it contains a dense mass of euspermatozoa
orienled with the heads towards the epithelium
of the duct and a mass of slender parasperma¬
tozoa in the center of the duct. The most distal
tenth of the duct contains mainly paraspermato¬
zoa (Fig. 9, sv). The sperm duct opens in the very

Source : MNHN, Paris

126

ANDERS WARÈN & PHILIPPE BOUCHET

proximal part of the palliai oviduct, close to the
openings of the receptaculum seminis. No ob-
vious connection to the sperm gutter on the pénis
(Fig. 8, seg) was found, unless a ciliated furrow
along the ventral part of the free lobe of the
palliai oviduct has such a function.

The female System consists of an open, glan-
dular palliai oviduct with six séminal réceptacles
(Fig. 8, r.se) opening via separate ducts to the
very proximal part of the oviduct (Fig. 8, pod).
The latter is presumably not fully formed.

The palliai oviduct is open (Figs 7. 26), except
its most distal and proximal parts (Fig. 3, 24, 8,
pod) which form wedge-shaped caeca in the
palliai skirt. The posterior caecum is continued
by a short blindly ending duct, to which the
séminal réceptacles open. via individual ducts.
Anteriorly to these opens vas deferens. Along the
broad edge of the free lobe of the oviduct runs a
strongly ciliated gutter (Figs 7, 26, If) which
starts at the very first part of the ventral slil and

continues anteriorly to shortly after where the
oviduct closes anteriorly. Therc the free lobe of
the gutter continues and transforms into a short
(tongue-like?) skinfold.

The proximal 1/3 of the oviduct is lined by a
low epithelium of several types of gland cells.
After this part a tall (40-50 pm) dark greyish
violet staining epithelium appears in the dorsal
part of the oviduct. This epithelium which is
interpreted as an albumen gland (Fig. 24, ag)
then spreads ventrally so it covers most of the
inside, except a narrow zone at the level of the
sût. The low epithelium spreads again, at the
beginning of the anterior caecum (Fig. 24), where
it covers the mid 1/3 of the heighl of the caecum.
The lower 1/3 is covered by tall (40-50 pm)
mucus producing epithelium (Fig. 24, mu), which
is continuous with the much lower but otherwise
identical epithelium of the corner of the palliai
cavity around the anterior part of the slit of the
oviduct.

Discussion

The capulids and trichotropids are presently
(Boss, 1982; Ponder & Warén, 1988 [as Capu-
lidae]) placed in the Calyptraeoidea, but there is
evidence from sperm morphology (J. Healy,
pers. comm.) that they actually are most closely
related to the Tonnoidea, a view we share. This
is also supported by further facts mentioned in
« Relationships ».

Furthermore, in the forthcoming discussion we
fully accept the view of Ponder & Warén (1988)
that the Capulidae and Trichotropidae are sy-
nonyms and use the name Capulidae to include
also the trichotropids.

Scarcity of material and validity of rcsults

The anatomical description is severely hampe-
red by the fact that very little material has been
available, and most of the information is derived
from a single specimen of Z. tropidophora. Serial
sections of the head-foot of H. gulerita showed
some différences in the structure of the oesopha-
gus and minor différences in the shape of the
ganglia of the central nervous System. The exter-
nal morphology is, however, quile similar as far

as could be observed from rehydrated bodies.
There were différences in the shape of the pénis
and the foot. but not very great and the latter
can to some extern be due to différences in
fixation and a resuit of the drying and rehydra¬
tion. There seem, however, to be présent two
types of penes; with or without an anterior lobe.

Larval shell

The morphology of the larval shell, with
distinct protoconch I and II présent in almost ail
species (not in H. japonica) indicates that they
hâve planktotrophic development. This is further
confirmed by the egg-capsule of //. carinata with
embryos of a size of 0.16 mm. which corresponds
well with the size of protoconch I in that species.

The larval shell is quite similar to some species
of Capulidae, see Figs 48-49. A few capulids are
known to hâve so-called echinospira larvae (Co-
pulus: Fretter & Graiiam, 1962; Trichosirius:
Pilkington, 1974, 1976). We hâve examined
protoconchs of several species known to hâve
echinospira larvae, from ail families where this
type of larval development is known and found
that they conchologically are characterized by

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

127

Figs. 28-34. Jaws and opcrcula of Haloceratidac. 28-30, Zygoceras iropidophora. - 28. operculum. hcighl 4.6 mm.
29-30, jaw, scale lincs 0.1 and 0.01 mm. 31, Haloceras cingulata, operculum. height 1.9 mm. 32, II. phaeocephala,
operculum, heighl 1.13 mm. 33, H. carinata, operculum. heighl 0.8 mm. 34, H. japonica. operculum. height 1.84

Source : MNHN, Paris

128

ANDERS WARÉN & PHILIPPE BOUCHET

Figs. 35-43. Radulac of Haloceratidae. 35, Zygoceras tropidophora. paralypc. 36. Haloceras carinata. Cancap stn
2.155. 37-38, H. tricarinata, Benthfdi stn 87. 39, H. tricarinata. " Discovery" sln 10141. 40 , H. japonica olï

Oregon. 41, H. cingulata. usnm 52077. 42. //. cingulata. USNM 94898. 43. H. pluieocepliala paralvnc Sc il,- il.,.

25 pm (36, 39. 40. 42) and 50 pm (35. 37. 38. 41, 43). es

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

129

the absence of a sharp distinction of protoconch
I and II. Judging from the presence of such a
distinct démarcation in the species of Haloceras,
we believe that the similarity to some capulids
cannot be used to infer that Haloceras has such
echinospira larvae.

Teleoconch

The more or less planispiral teleoconch. with a
large, irregularly shaped aperture and rapidly
increasing diameter of the whorls hints towards a
sedentary life probably on rocky bottoms or on
some other solid substrate.

Anatomical details

The shape of the front part of the foot was
always too distorted, to be used for comparisons
between the species. It does. however, imply a
sedentary mode of life. The division of the foot
into a rear part that contracts to a solid lump
and an anterior extensile part which contracts
more folded can be seen in many trichotropids
(Graham, 1954; Yonge, 1962; Warén unpubl.)
and is especially obvious in Capulus. Yonge
(1962) described the behaviour of Trichotropis
cancellata, which has a morphologically similar
foot. Specimens of that species sit attached by
the rear part of the foot, filtering sea-water with
the gill. When crawling the anterior part of the
foot can be extended far ahead. In Haloceras the
anterior part of the foot is evidently not used as
in Capulus , for holding and protecting the egg
mass (Thorson 1965), since an egg mass with
developing embryos was found in the palliai
cavity of one specimen, which possessed this kind
of foot.

The anterior pedal gland, the glandular layer
on the propodium and on the front part of the
mesopodium are unusually thick and certainly
hâve some functional importance in connection
with a sedentary life.

The very strongly contracted proboscis sheath.
the folded oesophagus, the long, coiled and
folded buccal and pedal connectives as well as
the length of other nerves emerging from the
cérébral ganglia. indieate that the proboscis can
be extended considerably and that probably the
cérébral ganglia move forwards during that pro-
cess.

It is not certain how the palliai parts of the
gonoduct function. It is obvious that (the own)
sperm is transported to the initial part of the
palliai oviduct by the vas deferens which also
functions as a séminal vesicle. But after that
there exist two possible ways: Either the sperm
continue forwards in the palliai oviduct. or it
leaves the palliai oviduct and uses the gutter
along the edge of the free lobe of the oviduct.
This gutter ends on a skin-fold on the outside of
the lobe, at the same level as the pénis. The
skin-fold could then hâve a function to transfer
the sperm to the pénis since there seems to be no
sperm gutter across the neck. This explanation,
however, is not likely because the sperm gutter of
the oviduct must be used for transport of sperm
from another male at copulation, unless sperm
can be transported both ways.

The absence of a bursa copulatrix can hardly
be explained by the fact that the specimen is not
a mature female, because, from a fertilization
point of view, it actually is mature since the
séminal réceptacles were full of sperm.

Biology

Very little identifiable stomach and rectal
contents were found. A foraminiferan was found
in a single specimen of Haloceras carinata.
Several other species had some unidentifiable
organic material, but there were no remains of
planktonic organisms or minerai particles. This
excludes filter feeding or grazing the détritus
cover of the boltom. since every such gastropod
we hâve examined, invariably has the stomach
and intestine filled with at least partly identifiable
matter. Such a feeding is also unlikely in species
with a well developed proboscis.

An active and predatory way of life is. howe¬
ver. unlikely because of the morphology of the
foot, which suggesls that this animal is adapted
for a sedentary life.

The presence of unusually large eyes in a
gastropod group almost exclusively living in the
deep sea is somewhal surprising. but the eyes are
not necessarily used during the benthic life. Thcy
may be a réminiscence from the planktonic life of
the larva. Bouchet & Warén (1986) found that
almost no deep-sea eulimids with lecithotrophic
development hâve eyes. while species with plank-
totrophic larvae usually hâve eyes. In some

Source : MNHN, Paris

130 ANDERS WARÊN &

species, however. the eyes are lost soon after
seulement.

The presence of a pénis in every specimen, also
those with unquestionably female features (egg
mass in palliai cavity. developing embryos in
oviduct) indicates that lhe species are either
simultaneous or protandric hermaphrodites. The
lutter is supported by the histology of the testis
of Z. tropidophora . which was almost exclusively
male, but with some immature eggs présent.
Hermaphroditism is mainly known from proso-
branchs with a sedenlary or not very active mode
of life.

A conséquence of the specimen of Z. tropido-
phoru being in the process of sex change is that it
may be far from full grown. since most protan-
drous hermaphroditic prosobranchs change sex
around or below half maximum size (e.g. Tricho-
tropis : Yonge, 1962; Eulima: Warén, 1984; Ca-
pulus: Thorson, 1965 and own observations).

Another remarkable feature in the biology ot
the species of Haloceratidae is their apparent
rarity. We know a total of 20 species from 103
specimens and shells. Only seven species are
known from more than five specimens, although
several occur in comparatively well investigaled
areas. Despite this rarity it is obvious that when
one species is présent, there is an unexpectedly
high probability to fmd at least one more species;

H. tricarinata, laxa, cingulata and carinata were
ail found at Seamount stn CP 30, SW of
Portugal in 2000m;

Haloceras japonica and millestriata were descri-
bed from the same dredge haul off S Japan,
1300 m;

H. laxa and carinata at “ Porcupine " Expédition
1870 stn 16. off Portugal in 1900 m;

H. aff. laxa and II. cingulata at " Thalassa " stn
Z435 in the northem part of the Bay of Biscay
in 1000m;

H. spinosa, phaeocephala and Haloceras sp. 1 at
" Kapala " stn 80-20-10, off New South Wales
in 1100 m;

H. biocalae and a damagcd undetermined species
al Biocal stn DW 48, off Southern New
Caledonia in 800 m;

H. cingulata and H. aff. laxa at Biogas stn
CP 37, Bay of Biscay in 2200 m.

Considering that this material cornes from
more than 2000 dredge hauls in the depth range

PHILIPPE BOUCHET

400-4000m, it is obvious that the catches were
not randomly distributed, but we do not know
any uniting factor, except that these stations are
sometimes unusually rich also in other gastro-
pods.

To summarize the biological features of the
two généra, we gel a group of not very active
deep-sea gastropods usually with planktotrophic
development, and some unknown, very spécial
requirements on the biotope. The species are
probaly predators.

Relationships

The anatomical and biological features outli-
ned above and in the description clearly indicate
that Haloceras and Zygoceras belong to the
Caenogastropoda, more precisely the Neolaenio-
glossa:

Taenioglossate radula; not taenioglossate in

Neogastropoda.

A simple oesophagus; with a valve of Leiblein

or similar structures in Neogastropoda.

Both zygo- and dialyneury; none of these in

Archaeotaenioglossa.

Protandrous hermaphoditism présent; very

rare in Neogastropoda.

There is. however. no neotaenioglossate family
in which Haloceras and Zygoceras can be inclu-
ded.

Several features point in different directions:

The larval shcll of the species with assumed
planktotrophic development shows the greatest
similarity to the genus Benthonella (Rissoidae,
Figs 46-47: B. tenella (Jeffreys, 1867); see also
Ponder, 1985; Bouchet, 1976). The distinct
coloration, size, and sculpture of protoconch 1
and II are quite similar. Also certain capulids, for
example Tricliotropis crassicostata Melvill, 1912,
hâve a similar larval shell, but it is smaller and
more planispiral (Figs 48-49).

The teleoconch does not resemble any other
group in a way that makes it possible to include
it there, although there is some resemblance to
the Capulidae, in that the shell is slightly irregu-
larly shaped, paucispiral with rapidly increasing
diameter, often has an irregular, rough sculpture
and has a well developed periostracum. These
features may, however, be convergences caused
by a sedentary life.

The foot resembles thaï of many Tonnoidea

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERAT1DAF.

131

Table 1. - Comparison of Haloceras, Capulidac and Tonnoidea.

CHARACTER

LAUBIERINIDAE *

CAPULIDAE

Haloceras

Foot wilh latéral furrow

yes

yes

yes

Posterior pcdal gland

?

poorly developed

présent

Hypobranchial gland

thin

thick

thin

Anterior alimentary canal

pleurembolic proboseis

very short pleurembolic
proboseis '

pleurembolic proboseis

Lowcr lip

unmodified

pseudoproboscis

unmodified

Jaws

présent

absent

présent

Salivary duels

long and slender

short

long and slender

Salivary ducts pass nerve ring

yes

no

yes and no

Accessory salivary glands

présent

absent

absent

Oesophageal gland

présent

absent

absent

Digestive gland openings

•distant from each other

distant fom cach other

close together

Crystalline style

•absent

présent

absent ?

Osphradium

monopectinate/bipectinate •

monopcctinate

bipectinate

Suboesophageal ganglion

with dialyneury

with or without dialyneury

with dialyneury

Supraocsophageal ganglion

?. with zygoneury *

with zygoneury

with zygoneury

Pedal ganglia

no accessory ganglia. ni
cross connective

o with accessory ganglia. no
cross connective

with accessory ganglia,
cross connective

Larval type

normal veligers

echinospira

normal veligers

Protoconch I distinct from II

yes

no

yes

Spcrm duel on head-foot

•open or closcd

open

not présent ?

Séminal réceptacle

•posterior, one or many

posterior. one

posterior. many

Bursa copulatrix

•anterior

absent

absent

Palliai oviduct

closed

open

open

Sperm gutter on palliai
oviduct

?no

no

yes

Gonopcricardial duct

unknown

no

no

Mode of feeding

•predatory

filterfeeders

predatory ?

An asterisk (*) indicatcs Ihai lhe character is noi known in the Laubierinidae and is exirapolated from other tonnoidean
familics. Anatomical details from Bouvier ( 1889). Df.i.i & Ponder (1964). Lacaze-Duthiers ( 1872. 1901). Warén & Bouchet
(1990). Yonge (1962).

1. Capulus ungaricus as well as other capulids do hâve a very short pleurembolic proboseis. as can bc seen froin the skin
fold bctween the tentaclcs. The pseudoproboscis actually is a drawn out part of the proboseis sheath (unpublishcd).

and the Capulidae in having a demarcated.
rough zone along and above lhe edge of the sole.
This is a rare character among gastropods and
has rarely been noticed, but we hâve seen it in
Aporrhaidae and Warén & Ponder (in press)
reported it in the Loxonematoidea.

The foot also resembles many capulids (espe-
cially Capulus), ail calyptraeids and some vani-
korids in being divided into two functionally
different parts. This is probably also functionally
evolved since it also occurs in Amathinidae
(Pyramidelloidea; Ponder. 1987). a totally unre-
lated group.

The operculum shows clear similarities to ail

capulids (Warén unpublished) and many ton-
noids (Warén & Bouchet, 1990) in having the
larval operculum remaining and clearly set off
from the postlarval part when young and then
losing the old part by some kind of érosion.

The alimentary System is very similar to that of
the Laubierinidae and some other of the less
modified Tonnoidea (see Warén & Bouchet.
1990). but differs in not having the complex and
large salivary glands présent in ail tonnoids
(except the Ficidae). Neither does it hâve a well
developed oesophagal gland.

The radula is quite similar to that of many of
the advanced mesogastropods, except some of

Source : MNHN, Paris

132

ANDF.RS WARÊN & PHILIPPE BOUCHET

Figs. 44-49. Cillina. Benthonelta, Tricholropis. 44-45. Cilhna globosa. syntypc usnm 186381. hcighl 2.8 mm. 46-47,
Benihonella lenella, Biomidi: 2 stn 6. 39°40'N, 04°57’E, 2610 m. hcighl 4.4 mm. 48-49. Triclwiropis crassicosiaia
"Marion-Dufresne" cruise 32, sln DC10. 21"I3'S, 55”52’E, 930-980 m. hcighl 5.6 mm. Scalc lincs 200 jim.

the more modified groups (e.g. Lamellaria, Cy-
praea, Pedicularia, etc.), but it is especially
similar to that of the two recently described
tonnoidean groups Laubierinidae and Pisanianu-
rinae (Ranellidae) (see Warén & Bouchet,
1990).

The anatomy and ontogeny of the reproduc¬
tive System is very similar to the Capulidae.
Direct comparison with sectioned specimens of
Capulus hungaricus, in the sexual phase when
oocyte production just has started showed a
similar construction of the oviduct. except that
Capulus has only a single séminal réceptacle, a
less developed ciliated furrow and a shorter
ventral sût of the oviduct.

On account of these characters we feel
confident that a position in the vicinity of the
Tonnoidea and the Capulidae is quite a realistic
assumption, and we hâve tabulated a number of
characters for the least modified tonnoidean
family and the Capulidae in Table I.

The larval development of the Capulidae, via
an echinospira larva is a character shared with
the Lamellariidae, Velutinidae, Triviidae and
Eratoidae (we do not consider the status of these
taxa, and the larva of Pseudosacculidae is not
known). Whether it is a homologous character
and its significance for these taxa being mono-
phyletic was discussed by Fretter & CJraham
( 1962:627). They concluded that this was the

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

133

case, based on Graham's (1954) comparative
study of Capulus, Tricholropis and Calyptraea,
but maintained that Capulus is more closely
related to Calyptraeoidea than to Lamellarioi-
dea. Since then Pilkington (1974. 1976) has
shown that also species of the old family Tricho-
tropidae hâve echinospira larvae, which supports
Fretter & Graham's ideas about Trichotropi-
dae as an ancestral group of the species posses-
sing echinospira larvae, the « Echinospiracea »
(modified to « Echinospirida » by Golikov &
Starobogatov, 1975). This (inding also contri-
buted to Ponder & Warén's (1988) view that the
Trichotropidae and Capulidae should be consi-
dered one family.

To our ideas about the Capulidae not being
related to the Calyptraeoidea, but to the groups
possessing an echinospira-larva should be added
the fact that ail taxa within the Calyptraeoidea
hâve a normal snout, while the Capulidae as well
as the Lamellarioidea hâve a proboscis (not to be
confused with the so-called pseudo-proboscis of
the Capulidae, which is an additional process
from the proboscis sheath).

If the echinospira larvae of the groups are
homologous (as we believe) this certainly subs-
tantiates the use of that type of larvae for
classification and makes it practical to maintain
them as a group, without mixing them with
species without that kind of larval development
(unless it is lecithotrophic).

We thus get a situation with a caenogastropod
lineage which splits into the Tonnoidea and the
Echinospirida (disregarding Hennigian principles
about ranks). The Laubierinidae and the Capu¬
lidae respectively are the most primitive members
of these two branches. The problem then is:
Where do Haloceras-Zygoceras join the « Y »?

There are actually no characters of the Echi¬
nospirida and Tonnoidea which are more primi¬
tive than in Haloceras-Zygoceras, but there are
characters in Haloceras and Zygoceras which are
more primitive, e.g. the closely placed openings
of the digestive glands and the accessory pedal
ganglia with a cross connection. The normal
veliger larva is more primitive than the echinos¬
pira larva (unless one assumes the normal veliger
to be secondary). This should support a position
either on the basal limb of the « Y » or on the
branch leading to the Tonnoidea, or on the
branch leading to the Echinospirida, before the
development of the echinospira larva took place.
The anatomical information available is, howe-
ver, hardly detailed enough to allow a clear
answer to where to place the two généra.

Whatever the relationships may be. Haloceras
and Zygoceras do not fit in any of the families
mentioned. why we suggest a new family for
them. We consider it related to but more primi¬
tive than the superfamily Tonnoidea and the
suborder Echinospirida.

Family HALOCERATIDAE fam. nov.

Diagnosis. Shell disc-shaped to fairly tall-
spired, with 1.5-3 teleoconch whorls covered by a
well developed, often fringed periostracum.
Sculpture usually spiral ribs and sinuous growth-
lines. Aperture rounded to depressed, without
siphonal canal. Umbilicus usually broad and
deep, occasionally closed. Protoconch 1 with
numerous small short raised ridges, often encir-
cling small round surfaces and usually aggrega-
ted to form 4-7 spiral ribs which are more well
defined towards periphery. Sometimes micro-
ridges are confined to spiral ribs and leave rest of
shell smooth. Protoconch II with 1-3 whorls, 2-3

spiral keels close to periphery. sometimes also
with pustules and other sculpture on last whorl.

Soft parts with bipectinate osphradium. pleu-
rembolic proboscis, simple salivary glands with
duels which may or may not pass nerve-ring.
fairly concentrated nervous System with dialy-
neury between supra- and cérébral, zygoneury
between suboesophageal and cérébral ganglia.
Two pairs of accessory pedal ganglia, posterior
pair with commissure. Foot with anterior and
posterior pedal glands and latéral furrow above
edge of sole. Protandrous or simultaneous her¬
maphrodites.

Source : MNHN, Paris

134

ANDERS WARÉN

PHILIPPE BOUCHET

SYSTEMATIC ACCOUNT
Genus HALOCERAS Dali, 1889

Separatista f Haloceras) Dali, 1889: 277. Type species:

Cithna cingulata Verrill, 1884, by monotypy.
Solariella (Micropiliscus) Dali, 1927:130. (New sy-

nonym). Type species: S.(M.) constricta Dali, 1927,

by original désignation.

Revised diagnosis. — Haloceratids with (in
species with planktotrophic development) mullis-
piral brown globular protoconch. Protoconch I
with 4-8 spiral cords on a background of nume-
rous anastomosing riblels and pustules. Proto¬
conch II with 3 spiral keels, of which two remain
uncovered above suture by successive whorl.
Teleoconch trochiform to lenticular, umbilicus
narrow to wide, whorls with 1 or 2 strong spiral
keels. Soft parts with the characters of the
family.

Remarks. — The history of the name has been
summarized in the introduction. For a compari-
son with Zygote ras, see that name.

Haloceras acrocomata sp. nov.

Figs 88. 93-94

Type material. Holotype in ams cl46278.

Type locality. “ Kimbla ” 1977, stn 3,
23°33.7'S, 152°37'E, 348-339 m, E of Lady
Musgrave Id, Queensland, Australia.

Material examined. Australia. “ Kimbla " 1977,
stn 3, E of Lady Musgrave Island, Queensland.
23°33.7'S, 152“37'E, 348-339 m: holotype (AMS
C146278). — Stn 22, ofT NW Island, Queensland,
23"15'S, 152°24'E, 284 m: 1 sh. (AMS C150187).
1984, stn 15, E of Lady Musgrave Island, Queensland,
23°52'N, 152°42'E, 296 m: 1 sh., 1 juv. sh. (AMS
Cl 47272).

Description. — Shell small, thin, solid,
consisting of 2 protoconch and 1.75 teleoconch
whorls. Protoconch rather fiat, almost planispi-
rally coiled. Protoconch I, diameter 240 pm,
sculptured by 6 strong spiral cords, continuous
from nucléus to protoconch II, except for a small
area near transition between protoconch I and
II, which is occupied by granules. Protoconch II

with 1.4 whorls. sculptured by two strong keels,
a third keel hidden at suture, visible only in
young postlarvae. Between keels, on later part of
protoconch II small granules; rest of larval shell
smooth or with occasional traces of periostracal
spirals. Protoconch/teleoconch transition sharp.
Teleoconch whorls turreted, broadly conical,
with two strong keels giving whorls a very
angular profile; no othcr spiral sculpture. Dis¬
crète axial sculpture made stronger by periostra¬
cal lamellae, best visible at periphery, where
forming projecting scales between keels. Umbili¬
cus broad, encircled by one strong thread. Aper-
ture broadly quadrangular; lip sharp, thin, only
very slightly reflected on columellar side. Colour
of shell very light amber white, with 2 carinae
and umbilical thread distinctly darker; proto¬
conch light brown.

Dimensions : Height 2.07 mm, breadth 3.06 mm;
aperture height 1.14 mm, breadth 1.26 mm.

Larval shell measurements:

diameter no. whorls

(t*m) (prot. Il)

hololype 940 1.4

ams 147279 1000 |,6

ams 142679 980 |.45

ams 150187 960 |.5

Remarks. One shell (ams 150187) has a
proportionally higher spire and narrower umbi¬
licus (the two characters are linked), but other-
wise identical protoconch and general morpho-
logy.

The two strong keels on the teleoconch of H.
acrocomata make it resemble IL exquisila and H.
cingulata. It differs from H. exquisita by its
low-spired protoconch with fewer whorls, by
teleoconch whorls which increase more rapidly in
diameter, and ils broader umbilicus. H. cingulata
reaches a much larger size at the same number of
postlarval whorls, and has a narrower umbilicus.

Etymoi.ogy. From the Greek akros, at the
top, and comatas, hairy; to remind of the hairy
periostracum of the protoconch.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

135

Figs. 50-55. Genus Haloceras. 50-52, II. tricarinata, Benthedi stn 87. diamcter 14.0min. 53-55, H. cingulaia ,

Biogas stn CP37, diametcr 13.8mm.

Source : MNHN, Paris

136

ANDERS WARÉN

PHILIPPE BOUCHET

Figs. 56-61. Genus Haloceras. 56, H. mediocostata. Monaco stn 703. heighl 3.2mm. 57, //. mediocostata, lectotype,

height 3.5mm. 58, H. carinata, holotypc of Solariella constricla, heighl 3.3mm. 59-60, //. carinata. Balgim stn

CP92. heighl 2.65mm. diamclcr 2.35mm. 61, H. carinata. Cancap stn 2.155. height 1.6mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

137

Haloceras carinata (Jeffreys, 1883)
Figs 33, 36, 58-61, 105-108

Cithna carinata JelTreys. 1883: 111, pl. 20, fig. 9.
Solariella (Micropiliscus) constricta Dali, 1927: 130.

(New synonym).

Solariella (Micropiliscus) constricta- Quinn 1979:43.

figs 67. 68.

Type material. — C. carinata: Holotype
BMNH 1885.11.5.1623; S. constricta : 11 syntypes,
usnm 108414a.b.

Type localities. C. carinata : “ Porcupine "
1870, stn 16, 39°55'N, 09°56'W. 1810m, olT
Portugal; S. constricta : “ Albatross ” stn 2415,
30°44'N, 79°26'W, 805m, off Florida.

Material examined. Eastern Atlantic. " Por¬
cupine ", stn 16, off Portugal. 39°55'N, 09°56'W. 1810
m: 1 sh., holotype.

Seamount, stn CP30, Gorringe Bank, 36°44'N,
11'23'W. 1940-2075m: 1 sh., 2 frgms (mnhn). Stn
DW116, Galicia Bank. 42°52'N, 1 l'SI'W, 985-1000m:
3 shs (SMNH).

Balgim, stn CP92. Ibero-Moroccan Gulf,
34°24.3'N, 07°30.3'W, 1182m: 1 sh. (mnhn).

Incal, stn DS3. W of the British Isles, 57°27'N,
11°03'W, 609-619m: 1 larva (lost).

Cancap. stn 2.067. Canary Islands. 27°58'N,
14°12'W, 1820m: 1 sh. (rmnh). Stn 2.155. Canary
Islands. 27°35'N, 17°59'W. 700m: 1 spm.. 3 shs
(rmnh).

Western Atlantic (Florida). “ Albatross ", stn 2415,
30°44'N, 79°26'W. 805 m. 11 shs (syntypes of S.
constricta Dali, 1927).

Description. Shell small. solid, conical,
with 2.5 protoconch and up to 2.75 teleoconch
whorls. Protoconch globular, multispiral. Proto¬
conch I diameter 200 pm, with sculpture of
star-shaped little knobs forming a réticulation,
and 5 spiral cords formed by coalescing such
knobs. Protoconch 11 with two spiral keels well
visible above suture, third keel hidden by subsé¬
quent whorl. Sculpture of granules appear in
subsutural zone after first whorl of protoconch
II, then gradually expends over most of proto¬
conch surface. 3-6 prosocline axial lamellae,
situated on last part of protoconch II, just
behind perislome. Protoconch lip expanded. Pro-
toconch/teleoconch discontinuity very abrupt.
Teleoconch trochoid with regularly convex
whorls and very deep suture. Sculpture consis-
ting mainly of spiral cords, two of which are

sometimes stronger, forming weak spiral keels.
About 25 spiral cords on penultimate whorl of
large specimens. Base and umbilicus sculplured
by similar cords. Border of umbilicus not marked
by cord. Spiral cords cross much weaker incré¬
mental lines. Above shoulder intersection of
spiral cords and incrémental lines produces a
kind of malleated surface, more distinct after
coating for SEM. Base convex. umbilicus open
and broad. Aperture rounded. peristome almost
continuous. Outer lip thin. sharp. slightly reflec-
ted over umbilicus. Colour of protoconch brown,
teleoconch dirty white.

Dimensions of shells: usnm 108414. height 3.24
mm. breadth 3.0 mm; aperture height 1.20 mm.
breadth 1.45 mm. Balgim stn CP92, height 2.62
mm. breadth 2.28 mm; aperture height 1.0 mm.
breadth 1.10 mm .

Larva! shell measurements:

diameter

(gm)

holotype carinata 840

usnm 108414 710

usnm 108414 720

usnm 108414 700

USNM 108414 770

USNM 108414 740

Cancap 2.067 980

Cancap 2.155 840

Cancap 2.155 730

Cancap 2.155 840

Cancap 2.155 890

Balgim CP 92 840

Seamount CP 30 920

Seamount DW 116 840

Seamount DW 116 900

Seamount DW 116 900

no. whorls
(prot. II)
1.8
1.9
1.8

1.7
1.9

1.8
1.75
1.9

1.7
1.75

1.8
1.9
1.9
1.8
1.9
1.75

Remarks. Warén (1980: 20). misled by the
similarities in the protoconch, commented thaï
the holotype of C. carinata is: « A monstrosity of
Benthonella tenella (Jeffreys) ».

Quinn (1979) listed as Solariella constricta ( =
carinata) three other lots which here are descri-
bed as Haloceras trichotropoides. Indeed, the two
species hâve rather similar teleoconchs, although
//. trichotropoides can be recognized by its
coarser sculpture, with fewer and stronger spiral
cords. They can also be separated by H. tricho¬
tropoides having a flatter larval shell with fewer
whorls.

Source : MNHN, Paris

138

ANDERS WARÉN & PHILIPPE BOUCHET

H. carinata also resembles H. mediocostata.
That species has a larger protoconch (diameter
1220 pm vs. 700-920 pm in carinata), and a
broader teleoconch sculptured by stronger spiral
keels.

Haloceras cingulata (Verrill, 1884)

Figs 31. 41-42. 53-55, 113-115

Cithna cingulata Verrill, 1884: 184, pl. 32, fig. 7.

Type material. Holotype usnm 38101.

Type locality. — “ Albatross ” stn 2076,
4P13'N, 66°01'W, 1658m, Georges Bank. S of
Nova Scotia.

Material examiner. — Western Atlantic. “ Alba¬
tross ”, stn 2043, S of Nova Scotia. 39°49'N, 68°29'W,
2685 m: 1 spm., paratype (usnm 38104). Stn 2076,
Georges Bank. S of Nova Scotia. 41°13'N. 66°01'W.
1658 m: 1 sh., holotype (usnm 38101). Stn 2084, S
of Nova Scotia. 40°17'N, 67°05'W, 2361m, I sh..
paratype (usnm 38105). — Stn 2571. S of Nova Scotia,
40°09'N, 67°09'W, 2480m: I spm. (usnm 52077). Stn
2733, ofl North Carolina. 37'26'N, 73"43'W. 1728m: 1
spm. (usnm 94898).

Eastern Atlantic. Biogas, stn CP 37, Bay of Biscay,
47°34'N, 08°40'W, 2175 m: I spm. (mnhn).

“ Thalassa ", Stn Z 435, Bay of Biscay, 48°40'N,
09°56'W. 1050 m: 1 sh. (mnhn).

Seamount, stn CP30, Gorringe Bank, 36°44'N,
H°23'W. 1940- 2075m: 7 juv. and frgms (mnhn).

30 on base, also in umbilicus. In addition to
incrémental lines, also a few coarse, strongly
prosocline ribs on shoulder of terminal part of
body-whorl. Umbilicus very broad. Aperture
rectangular. peristome almost continuous; outer
lip sharp, simple. Pariétal wall convex, strongly
reflecled towards umbilicus. Colour of proto¬
conch brown. teleoconch dirty white.

Dimensions of a fully grown shell: Height
8.1 mm, diameter 13.2 mm; aperture height 4.3
mm, breadth 7.7 mm.

Larval shell measurements:

diameter
(pm)

usnm 52077 1380

Seamount CP 30 1510

Seamount CP 30 1490

Seamount CP 30 1660

Seamount CP 30 1580

Biogas CP 37 1340

“ Thalassa " Z435 1300

no. whorls
(prol. Il)
2.2
2.2
2.25
2.3
2.1

2.1

Remarks. There is no species easily confu-
sed with H. cingulata. The two strong spiral keels
appear early on the teleoconch, making
identification ol very young specimens possible.
At comparable size only Zygoceras biocalae
bears some resemblance but lacks almost
completely a spiral sculpture and has a much
smaller umbilicus.

Distribution. — The temperate North Atlan¬
tic, between 36° and 49°N. in 1050-2685m (bath-
yal); amphiatlantic.

Description. — Shell large, solid, depressed,
consisting of 2.5 protoconch and up to 2.6
teleoconch whorls. Protoconch globular, multis-
piral. Protoconch I diameter 250 pm, sculpture
not very well preserved on available specimens,
but réticulation of anastomosing star-shaped
knobs présent beside a few (3-5 ?) spiral cords.
Protoconch II with two spiral keels apparent
above suture, third keel concealed by subséquent
whorl. Last half protoconch II whorl with dis¬
crète sculpture of tubercles in subsutural zone.
Protoconch/teleoconch transition very sharp. Te¬
leoconch with strongly keeled whorls, almost
rectangular in cross-section, adhering to prece-
ding whorl just below abapical keel, and concea-
ling suture in deep furrow. Sculpture consisting
of spiral cords and incrémental lines. 25 spiral
cords between suture and first keel on adapical
part of body whorl; 6 between keels; and about

Haloceras exquisita sp. nov.

Figs 83, 89-90

Type material. Holotype in mnhn.

Type locality. Biocal, stn DW66, 24°55'S
168°22'E, 505-515m, S of New Caledonia.

Material examined. New Caledonia. Biocal
stn DW 66. 24°55'S, I68°22'E, 505-515 m: holotype
(mnhn).

Description. Shell small, thin, fragile,
consisting of 2.5 protoconch and 1.5 teleoconch
whorls. Larval shell globular, multispiral. Proto¬
conch I diameter 215 pm, sculptured with rather
irregularly disposed knobs in nucléus, later ar-
ranged in spiral lines. Protoconch II with 2.1
whorls, diameter 870 pm; first whorl sculptured
by onc strong spiral keel and one row of
subsutural granules; later a second spiral keel
previously hidden by suture, and a sculpture of

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

139

small knobs and granules appears, between keels.
Surface of last half whorl of protoconch II
completeiy covered by such granules and knobs.
Teleoconch whorls turreted. wilh two very strong
keels at periphery giving a very angular profile to
whorls. No other spiral sculpture except for a
discrète thread bordering umbilicus. Upper part
of whorls with sculpture of prosocline axial ribs
extending from suture to adapical keel. forming
spiny intersections with keel. Axial ribs obsolète
between keels. Base smooth. Umbilicus narrow.
Aperture quadrangular. outer lip thin, sharp. a
little reflected over umbilicus. Colour of proto¬
conch brown. teleoconch white.

Dimensions : Height 1.80 mm, breadth 1.75
mm; aperture height 0.72 mm, breadth 0.77 mm.

Remarks. Haloceras exquisita has a very
distinctive protoconch and teleoconch morpho-
logy which should facilitate identification. The
protoconch is smaller, at a similar number of
whorls, than in any other haloceratid. The
general morphology and sculpture of the teleo¬
conch whorls resemble H. acrocomata, but the
whorls are not as depressed as in H. acrocomata.

Etymology. From the latin exquisitus , fine.

Haloceras galerita sp. nov.

Figs 66, 100-102

Type material. Holotype ams Cl 50227.

Type locality. — “ Kapala ”, stn K80.20.20,
33°35-37'S, 152°05'E. 1143-1106m, off Sydney,
New South Wales.

Material examined. Australia. " Kapala ", stn
K 80.20.20., off Sydney. 33°35'S, I52°05’E, 1143-1106
m: holotype (ams Cl50227).

Description. — Shell small. solid, trochoid,
consisting of 2.5 protoconch and 1.9 teleoconch
whorls. Protoconch I diameter 200 pm, sculpture
not very distinct but seemingly consisting of
coarse spiral cords. Protoconch II with 2 spiral
keels visible above suture, and a subsutural row
of granules. Also a few distinct incrémental ribs
just before larval peristome; rest of protoconch
smooth. Teleoconch whorls convex wilh rather
deep suture. Sculpture of spiral cords and strong

incrémental, almost lamellar, fines. Two spiral
cords form keels; abapical one. sharp and clearly
defining basal surface. About 8 spiral cords
above shoulder of body whorl, 6 at periphery
between keels, and about 18 on basal area,
extending into umbilical région. Incrémental fi¬
nes strongly curved, with most retracted point on
adapical keel. Umbilicus narrow. Aperture
ovoid. broader than high. Outer lip thin, simple,
a little reflected over umbilicus. Colour of pro¬
toconch brown, teleoconch white.

Dimensions: Height 2.6 mm, breadth 2.8 mm;
aperture height 1.2 mm, breadth 1.5 mm.

Larval shell measurements:

diameler no.whorls

(gm) (prot.II)

1080 2.0

Remarks. - Haloceras galerita resembles H.
mediocostata, but it has a stronger. coarser spiral
sculpture, with better defined spiral keels, the
umbilicus is even more narrow and the diameter
of the whorls increases less rapidly than in
mediocostata.

Etymology. - From the Latin galerum. a hat
or helmet; to remind of the large and conspi-
cuous protoconch.

Haloceras heliptyx sp. nov.

(Figs 63. 103)

Type material. — Holotype in mnhn.

Type locality. Biogeocal, stn DW289.
20°36'S. 167°00'E, 830-840 m, Loyalty Basin. E
of New Caledonia.

Material examined. Loyalty Basin (E of New
Caledonia). Biogeocal, stn DW 289, 20'36'S,
167°00'E, 830-840 m: holotype (mnhn).

Description. — Shell small, solid, trochoid
consisting of 2.75 teleoconch, and unknown
number of protoconch whorls (apex broken).
Protoconch probably globular; little more than
last whorl remaining, diameter 770 pm. Proto¬
conch 11 sculptured with two strong spiral
keels, one at periphery. one just above suture;
zone between two keels with numerous small

Source : MNHN, Paris

140

ANDERS WARÉN & PHILIPPE BOUCHET

pustules. Flexuous axial riblets above peripheral
keel on last part of protoconch II. Teleoconch
whorls very convex, with slightly channellcd
suture. Spiral sculpture stronger than axial sculp¬
ture, consisting of three strong keels and much
weaker spiral cords. Abapical spiral keel hidden
at suture on spire, apparent only on body whorl.
6 much finer cords on shoulder of whorls, one
between keels, and 10 on basal area. Axial
sculpture consisting of strongly prosocline cords,
not extending to basal area and forming little
nodules at intersections with spiral keels. Also
numerous second order incrémental ridges, ex¬
tending over base. Umbilicus open, smooth
inside. Aperture ovate, outer lip rather regularly
convex, columellar région slightly reflected over
umbilicus. Colour of protoconch brown, teleo¬
conch white.

Dimensions : Height 3.1 mm, breadth 2.9 mm;
aperture height 1.0 mm, breadth 1.25 mm.

Remarks. — H. heliplyx resembles H. galerila,
but differs by having stronger axial sculpture and
fewer and stronger spiral cords. The umbilicus of
H. galerita is almost closed and its protoconch II
has a bigger diameter, 1080 pm. There is also a
general resemblance to H. japonica , which has a
thinner shell with thick periostracum and roun-
ded aperture, and differs in its brooding repro¬
ductive biology.

Etymology. From the Greek hélix , spiral,
and ptyx , a fold; to remind of the strong spiral
keels of the shell.

Haloceras japonica Okutani, 1964
Figs 34, 40, 62. 64-65

Halocerus japonicus Okutani, 1964: 397, pl. 6, fig. 8.

Type material. Holotype and I paratype
in Tokyo University Muséum, RM 8824.

Type locality. 34°10'N, 140°05.5'E, 1230-
1350m, 26 miles off Miyaké Island, Japan.

Material examined. Japan. 26 miles off Miyaké
Island. 34°10'N, 140°05.5'E, 1230-1350 m: holotype, 1
paratype (Tokyo Univ. Mus. RM 8824).

Eastern Pacifie (Oregon). “ Yaquina ", stn OTB-188,
45°50'N. 125°14'W. 1580 m: 1 spm. (lacm 67-167.2).

Description. Teleoconch high, trochoid,
consisting of more than 2.5 whorls (corroded)
with thick, strongly adhering periostracum.
Whorls convex, suture moderately deep. Sculp¬
ture consisting of spiral cords, of which two form
keels at periphery. no spiral cords above shoul¬
der, 2 between keels and 5 on base. Axial
sculpture of periostracal lamellae. Umbilicus
broad, encircled by a broad but indistinct cord.
Aperture rounded, forming a slight angle al base
of columellar pillar. Outer lip thin, sharp. Colour
of shell chalky white, periostracum dark greenish
brown.

Dimensions of holotype : height 5.95 mm.
breadth 5.05 mm; diameter of the aperture 2.25
mm.

Remarks. Neither of the two known shells
hâve a well preserved protoconch. The dried
body of the specimen from off Oregon was found
to contain 9 brooded embryos (Fig. 64). The
shells of some of these embryos were extracted
(Fig. 65). They are white, 1030-1050 gm in
diameter, with a sculpture of coarse granules
extending half a whorl further than on proto¬
conch I of other haloceratids. The assignment of
the species to Haloceras rather than Zygoceras is
based on the general resemblance in the shape of
the teleoconch compared with other species of
Haloceras. primarily H. galerita and H. heliptyx.

Haloceras taxa (JelTreys, 1885)

Figs 74-75, 116-118

Seguenzia taxa Jeffreys, 1885: 44. pl.5, figs 4-4a.

Type material. Holotype bmnh

1885.11.5.2586.

Type locality. “ Porcupine " 1870, stn 16
39°55'N, 09°56'W, 1810m, off Portugal.

Material examined. Eastern Atlantic. “ p or .
cupine ", 1870, stn 16, off Portugal, 39°55'N, 09°56'W
1810 m: holotype (bmnh 1885. 11.5.2586).

Monaco, stn 624. Azores, 38*59'N, 28°18'W
2102m. I sh. (mom).

“ Sarsia", stn 7614, Bay of Biscay, 43°4V1\I
03°38'W, 1100m: I sh. (bmnh).

Seamount. stn CP 30, Gorringe Bank. 36°44'N
11°23'W. 1940-2075 m: 7 shs (mnhn).

Three additional shells are tentatively identified as
Haloceras aff. laxa:

“ Thalassa “ stn Y 378, off NW Snain. 41°34'N
09°16'W. 1000m: I sh. (mnhn). Stn Z 435, Bay of

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

141

Biscay, 48°40'N, 09°56'W. 1050m: 1 sh. (mnhn).

Biogas, stn CP 37, 47°34'N, 08°39'W, 2175m: 1 sh
(mnhn).

Description. — Shell small, solid, discoid,
consisting of 2.3 protoconch whorls and up to
1.4 teleoconch whorls. Protoconch globular,
multispiral. Protoconch I diameter 220 pm,
sculptured by combination of anastomosing star-
shaped knobs and spiral cords. Protoconch
II smooth, except two strong spiral keels and
one subsutural row of granules. Protoconch/te-
leoconch transition abrupt. Teleoconch whorls
convex, first rather regularly rounded in young
postlarva, then cross-section of whorl becomes
more and more depressed, ending up in a
horizontally ovoid cross-section of body-whorl,
with very deep suture. Sculpture consisting of
strong spiral cords and incrémental Unes. Cords
strongest at periphery and weaker or obsolète on
shoulder and base. Incrémental Unes very regular
and closely set, giving shell a shining appearance,
especially on shoulder. Umbilicus wide, not
encircled by cord. Aperture horizontally ovoid,
peristome continuons, only slightly adhering to
penultimate whorl. Inner Up strongly reflected
over umbilicus. Colour of protoconch I brown,
protoconch II much paler except at protoconch-
/teleoconch transition, teleoconch white.

Dimensions of largest shell: Height 2.1 mm,
diameter 3.4 mm; aperture height 0.9 mm,
breadth 1.9 mm.

Larval shell measurements :

diameter

(pm)

Seamount CP 30 1130

Seamount CP 30 1150

Seamount CP 30 1100

Seamount CP 30 1100

Seamount CP 30 1140

Seamount CP 30 1140

Monaco 624 1200

“ Sarsia " 7614 980

afT. taxa

“ Thalassa " Y378 1030

" Thalassa " Z435 1060

Biogas CP 37 1100

no.whorls

(prot.ll)

1.8

1.8

1.75

1.75

1.75

1.8

1.75

1.85

1.8

Remakks. The holotype of S. laxa is 4.2mm
high. it is badly broken and wom which makes

identification of the name very difficult. Al-
though the identification at family level is beyond
doubt, there are no specimens collected in the
NE Atlantic that match this fragment precisely.
It could be a fragment of a very large H. carinata
(but there is no specimen of H. carinata higher
than 3.3 mm known), or it could be a fragment
of the species we hâve used the name for (but it
has less depressed whorls than our shells of that
size). It is probable that when more NE Atlantic
specimens become available, it will be possible to
match this fragment more accurately, but it is
incredible that Jeffreys based a new species on
this wom fragment.

H. laxa can be identified from olher species by
the absence of well defined keels at the periphery
of teleoconch whorls, its depressed whorls. and
small adult size. H. cingulata is similarly depres¬
sed, but has strongly keeled whorls and a much
larger adult size. H. carinata also lacks well
defined spiral keels but it has a higher spire and
considerably smaller protoconch. H. rugosa,
from the SE Pacific, has an almost closed
umbilicus and a distinctive sculpture on the
adapical part of teleoconch whorls.

The shells referred to as H. aff. laxa differ in
having a stronger spiral sculpture, and more
regularly rounded, not depressed whorls. The
protoconch sculpture and measurements are ho-
wever indistinguishable from those in typical
laxa. Two of these shells originate from the
northern part of the Bay of Biscay. while the
more depressed specimens are from further
south. They may represent variation or belong to
a distinct species.

Haloceras sp. I
Figs 68-69, 70-71, 104

Material examined. New Caledonia. Biocai..
stn DW 70. 23-25'S, 167°53'E. 965 ni: 2 shs (mnhn).

Australia. " Kapala" stn 80-20-10. off Svdnev.
33'36'S, 152°05'E, 1105-1143m: 3 shs (ams C157269).

Description. — Shell trochoid, with a low
spire. Protoconch I, diameter 230 pm. not per-
fectly preserved but with réticulation of anasto¬
mosing knobs and spiral cords. Protoconch II
with 2 visible spiral keels, one well visible and
one at or just above suture. Subsutural zone
granulated. Teleoconch with up to 1.9 convex.

Source : MNHN, Paris

142

ANDERS WARÊN & PHILIPPE BOUCHET

regularly rounded whorls, sculptured with spiral
cords and incrémental lines. Two spiral cords
form indistinct keels at periphery. Umbilicus is
open.

Dimensions of largest sheli. Height 2.7 mm,
breadth 3.1 mm.

océan. Considering the vast distances (more than
20,000 km) separating the locations, we judge the
material insufficient to conclude if one, somew-
hat variable, or several, closely related species are
involved.

Larval sheli measurements :

diameter

(nm)

Biocal DW 70 1020
Biocal DW 70 1080
AMS 157269 900
ams 157269 920
ams 157269 900

no. whorls
(prot.II)
1.9
1.9
1.75
1.95
2.0

Haloceras mediocostata
(Dautzenberg & Fischer, 1896)
Figs 56-57, 97-99

Cithna carinata var.

Fischer. 1896: 449.
Cithna jeffrevsi var.

Fischer. 1897: 159.
Cithna carinata var.
1927: 118.

mediocostata Daulzenberg &
mediocostata Dautzenberg &
mediocostata — Dautzenberg

Remarks. — These shells are somewhat inter-
mediate between H. carinata and taxa. They hâve
a higher spire and a stronger spiral sculpture
than H. taxa, and a lower spire and bigger
protoconch than H. carinata. There is no doubt
however that Haloceras taxa and H. carinata are
two distinct species in the North Atlantic.

In fact these shells strongly resemble the NE
Atlantic shells we hâve called Haloceras aff. taxa
above, and may be conspecific with them. We
would probably hâve treated them as a single
species if they had originated from the same

Type material. Lectotype, here designated,
one sheli marked by Dautzenberg « type » in
mom (Fig. 57); paralectotype from the same
station in mnhn.

Type locai.ity. Monaco, stn 553, 37°43'N
25°05'W, 1385m, Azores.

Material examined. Fastern Allantic. Monaco
stn 553. Azores. 37°43'N. 25°05'W. 1385 m: Lectotynè
(mom): 1 paralectotype (mnhn). Stn 578, 38°26'N
26°3I'W, 1165m: I sh. Stn 703, 39'21’N 3|°06’w'
1360m: 2 shs. ‘ W '

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

143

Figs. 64-65. Haloceras japonica, off Oregon. 64,
Specimen wilh partly broken body whorl showing lhe
cmbryos brooded in ihc oviduct; scale line 1 mm.

65, Larval shell of thc cmbryos; scale line 0.25 mm.

Dautzenburg (1927) also listed material from stn
203, 683, 698 and 719. We hâve examined this material
in mom: the shell from stn 683 belongs to an undes-
cribed species of Zygoceras (see p. 159); others are
unrecognizable fragments, or hâve been destroyed by
acidic glass tubes.

suture slightly channeled but otherwise only
moderately deep. Sculpture of spiral cords and
incrémental lines. Two stronger cords form spiral
keels at periphery; about 8 spiral cords on
shoulder of body whorl, 3 at periphery between
keels; 12 on base, extending to umbilical région.
Umbilicus rather narrow, not encircled by cord.
Aperture rounded; outer lip thin, simple, inner
lip a Utile reflected over umbilicus. Colour of
shell white, protoconch brown.

Dimensions of a complété shell : Height 2.5 mm,
breadth 2.6 mm; aperture height 1.2 mm, breadth
1.3 mm.

Larval shell measurements:

diameter

(gm)

Monaco 553 1220

Monaco 553 1240

Monaco 578 1200

Monaco 703 1300

Monaco 703 1240

(prot.II)

2.1

11

2.0

2.1

2.0

Remarks. — The assignment of this species to
Cithna jeffreysi [= Benthonella tenella (Jeffreys,
1869); Rissoidae] by Dautzenberg & Fischer,
1897 was due to a printer's mistake (Dautzen¬
berg. 1927).

H. mediocostata bears some resemblance to the
keeled specimens of H. carinata (which lives in
the same part of the North Atlantic), but can be
separated by the size of the protoconch: diameter
700-920 gm (mean 817 pm) in H. carinata, vs.
1200-1300 pm (mean 1240 pm) in H. mediocos¬
tata-, H. carinata also has a higher spire.

Haloceras millestriata (Okutani, 1964)
Fig. 67

Description. Shell small, thin, fragile,
trochoid, consisting of 2.5 protoconch and up to
2.3 teleoconch whorls. Protoconch globular,
multispiral. Protoconch I, diameter 200 pm, with
a sculpture of spiral cords. Protoconch II with
two spiral keels (one at or just above suture) and
a subsutural row of granules. In last part of
protoconch II, just before larval peristome, a few
strong incrémental ribs; rest of protoconch
smooth. Teleoconch whorls regularly convex.

Torellia millestriata Okutani. 1964: 397, pl. 6, fig. 4.

Type material. Holotype in Tokyo Uni-
versity Muséum, rm 8822.

Type locality. — 34°10'N, 140°05.5'E, 1230-
1350m, 26 miles off Miyaké Island. Japan.

Material examined. — Japan. 26 miles off Miyaké
Island, 34°10'N, I40°05.5'E. 1230-1350 m: holotype
(Tokyo Univ. Mus., rm 8822).

Source : MNHN, Paris

Figs. 66-71. Genus Haloceras. 66, H. galerita , holotype, heighl 2.6mm. 67, H. milleslriala, holotype, height 3.3mm

68-69, Haloceras sp.l, ams C 157269. diameter 2.35mm. heighl 2.65mm. 70-71, Haloceras sp. I. Bloc ai. sln DW70

diamcter 2.9mm. heighl 2.5mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

145

Remarks. - The holotype and only known
shell of H. millesiriaiti has no larval shell. Our
generic assignment is based on the general mor-
phology of the teleoconch. The whorls are de-
pressed and the spiral sculpture forms two
indistinct keels at the periphery. Haloceras mil-
lestriala ressembles a little H. luxa and H.
rugosa, which both hâve a low spire, and regu-
larly rounded whorls. It is however easily reco-
gnizable by its even sculpture of strong spiral
lines, and moderalely open umbilicus.

Haloceras phaeocephala sp. nov.

Figs 32, 43. 81-82, 95-96

Type material. Holotype and 2 paratypes
(1 live taken) ams Cl 46202.

Type locality. “ Kapala" 1980, 33°36'S,
152°05'E, 1143-1106m, off Sydney, New South
Wales, Australia.

Materiai. examined. Australia. " Kapala" 1980, off
Sydney, 33°36'S, 152°03'E, 1143-1106 m: holotype, 2
paratypes (ams Cl46202). Off Sydney, 33°38'S,
I52°03'E, 924-896m: 1 sh. (ams Cl 50189). Off Wollon-
gong, New South Wales, 34°26'S. 15r27'E. 1200m: 2 shs
(ams C150188). NE of Batemans Bay, New South
Wales, 35'30'S, 150"52'E. 960-969m: I spm. (ams
Cl 50190).

Description. Shell small, solid, depressed
rissoiform, consisting of 2.3 protoconch and 2
teleoconch whorls with thick adhering periostra-
cum. Protoconch 1, diameter 230 pm. nucléus
with thin spiral threads which soon stop, leaving
rest of protoconch I smooth, except for 4 spiral
threads above suture. First whorl of protoconch
II sculptured by continuation of spiral threads
from protoconch I. Later sculpture of irregular,
raised scratches and spirally elongated knobs
develops and completely covers last protoconch
whorl. Protoconch/teleoconch discontinuity very
sharp. Teleoconch whorls convex, shouldered,
with moderately deep suture. Sculpture consists
of strong, broad axial ribs, thin incrémental
lamellae and spiral cords. 11 axial ribs on first
teleoconch whorl, and 15 on body whorl of
holotype. 5 strong spiral cords at periphery of
which adapical one forming a slightly spiny
shoulder at intersections with axial ribs. 2, later
on body-whorl 3, fainter spiral cords on shoul¬
der, 8 on base, of which one demarcates umbi¬
licus. Umbilical chink narrow. Ouler lip thin,
simple, with most projccting point at lower third;

inner lip slightly reflected and forming narrow
callus over umbilicus, a little protracted at base.
Colour of larval shell reddish brown. teleoconch
with yellowish white periostracum.

Dimensions: Height 3.33 mm, breadth 3.06 mm:
aperture height 1.74 mm, breadth 1.50 mm.

Larval shell measurements:

ams 146202
(holotype)
ams 146202
ams 146202
ams 150188
ams 150189
ams 150190

diameter

(gm)

1060

940

1000

1000

910

890

no.whorls

(prot.Il)

2.0

1.8

1.9

1.9

1.8

Remarks. — There is variation in the strength
of the sculpture. One of the shells (ams 150189)
is almost devoid of axial ribs, but transitional
specimens are présent, although sculptured spé¬
cimens appear to be more frequent. The sculp¬
ture, size and number of whorls of the proto¬
conch are the same in strongly and weakly
sculptured shells and they are undoubtedly
conspecific.

The protoconch of H. phaeocephala differs
from that of congeners in having a protoconch I
with spiral threads on a smooth background. a
character of Zygoceras. Protoconch 11 however is
characteristic for the genus.

Récognition of this distinctive little haloceratid
should be easy since it is the only représentative
of the family with a strongly sculptured. rissoi¬
form shell.

Etymology. — From the Greek pliaios.
brown, and kephale , head; to remind of the
brown coloured protoconch.

Haloceras rugosa sp. nov.

Figs 72-73, 91-92

Type material. Holotype lacm 2279.

Type locality. Allan Hancock Founda¬
tion, stn 482, 01°09'S, 90°36'W, 455m, Galapagos
Islands.

Material examined. Galapagos Islands. Allan
Hancock Foundation, stn 482, 01°09'S, 90°36'W,
455 m: holotype (lacm 2279).

Source : MNHN, Paris

146

ANDERS WARÈN & PHILIPPE BOUCHET

Figs 72-77. Généra Haloceras and Zygoceras. 72-73, Haloceras rugosa, holotypc. hcight 2.95mm, diameter 3.65mm.
74-75, H. taxa. Seamount sin CP30, height 2.1mm, diameler 3.4mm. 76-77, Zygoceras biocalae, holotypc, height

7.0mm, diameter 9.6mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

147

Description. Shell solid. globular. consis-
ting of 2.5 protoconch and 1.25 teleoconch
whorls. Protoconch I, diameter 200 pm, with
about 10 irregular spirals. interconnected by a
net of anastomosing short trabecles. Protoconch
Il with two spiral keels; one well exposed, one
partly hidden at suture; rest of protoconch
smooth except for one row of subsutural granu¬
les. Teleoconch whorls regularly convex. with
deep suture. Umbilicus open. narrow. Sculpture
above periphery of diverging, strongly opistho-
cline radiating ribs which intersect rather coarse,
prosocline incrémental ribs and produce irregu-
lar hammered appearance. 5-6 regular spiral
cords at and below periphery; another group of
similar spiral cords, more crowded towards basal
and umbilical région. Incrémental sculpture
weak and indistinct. Aperture simple, outer lip
sharp, thin. inner lip slightly reflected over
umbilicus. Protoconch brown. teleoconch white.

Dimensions of shell: Height 2.95 mm. diameter
3.65 mm; aperture height 1.80 mm, breadth 1.95
mm.

Larval shell measurements:

diameter no.whorls

(Hm) (prot.II)

1100 2.1

Remarks. H. rugosa is characterized by the
malleated appearence of the surface of the ada-
pical part of its whorls, with the strongly opis-
thocline ribs that cross prosocline incrémental
ribs. This character, in combination with the
small umbilicus and regularly rounded, unkeeled
whorl facilitâtes the identification.

Etymology. From the Latin rugosus , sch-
riveled, wrinkled; to remind of the teleoconch
sculpture.

Haloceras spinosa sp. nov.

Figs 80, 109-110

Type material. — Holotype ams cl57270.

Type locality. "Kapala", stn K80.20.10,
33°36'S. I52°03'E. 1106-1143m. off Sydney, New
South Wales, Australia.

Material examined. Australia. " Kapala ”, stn
K 79.20.14, off Broken Bay. New South Wales,

33°38'S. 152°06'E, 1033 m: 1 sh. (ams C146529). Stn
K 80.20.10, off Sydney. 33°36'S. 152'03'E. 1106-1143
ni: holotype (ams Cl 57270).

Lovaltv Basin (E New Caledonia). BlOGEOCAL, stn
CP 232, 21°34'S, 166‘27'E. 760-790 m: 2 shs (mnhn).

Description. — Shell small, solid, broadly
rissoiform, consisting of 1.7 protoconch and 2.7
teleoconch whorls, with thin, transparent, adhe-
ring periostracum. Protoconch I. diameter 290
pm, sculptured with about 10 rather fine spiral
threads. Shell corroded at transition protoconch
I/II. Protoconch II smooth and shiny except for
narrow subsutural zone of granules, and two
strong spiral keels at periphery. Protoconch
teleoconch transition abrupt, with peristome of
protoconch slightly flaring. Teleoconch whorls
convex with rather deep channelled suture.
Sculpture consisting of strong spiral cords and
weaker, lamellar axial ribs, forming small raised
spines when they intersect. 4 spiral cords on first
whorl, secondary spirals appear on body whorl
between adapical cords; also one cord encircling
slightly convex base, 4 other cords on base and 2
more indistinct ones in umbilicus. Channeled
appearance of suture results from it being lined
by two adjacent raised spiral cords. 19 prosocline
axial ribs on first teleoconch whorl, 32 on body
whorl. Between these, fine incrémental lines.
Umbilicus open, small. Aperture slightly depres-
sed; outer lip sharp, simple; inner lip slightly
reflected over umbilicus. Colour of larval shell
brown, teleoconch white.

Dimensions : Height 3.28 mm. breadth 2.88
mm; aperture height 1.12 mm, breadth 1.32 mm.

Larval shell measurements:

diameter no.whorls

(pm) (prot.II)

holotype 720 1.2

Biogeocal CP 232 675 1.3

BlOGEOCAL CP 232 690 1.3

Remarks. H. spinosa is immediately sepa-
rated from other haloceratids by the spiny gene¬
ral appearance.

Etymology. From the Latin spinosus ,
thorny; to remind of the teleoconch sculpture.

Source : MNHN, Paris

148

ANDERS WARÉN

PHILIPPE BOUCHET

Figs. 78-82. — Genus Haloceras. 78-79, //. Iricholropoides, holotypc. heighl 3.57 mm. diametcr 3.06min. 80, //. xpinosa,

holôtype. heighl 3.28mm. 81, //. phaeocephala, holotypc. heighl 3.33 mm. 82, II. phaeocephala, ams C 150189]

heighl 2.50mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NFW FAMILY HALOCERATIDAE

149

Haloceras tricarinata (Jeffreys, 1885)

Figs 37-39. 50-52. 119-120

Seguenzia tricarinata Jeffreys, 1885: 43. pl.5, fig.2.
Seguenzia tricarinata- Quinn 1983: 744.

Type material. Holotype bmnh 1885.11.
5.2589.

Type locality. — " Porcupine ” 1870, stn
17a, 39°39'N, 09°39'W. 1347m, W of Portugal.

consists of spiral cords and thin. raised, perios-
tracal incrémental lamellae. About 15 spiral
cords above periphery on early teleoconch, about
20 on later parts. Base of shell and umbilical
région with about 25 such spiral cords. Outer lip
thin, simple, angular at periphery: inner lip
sharp, simple, slightly reflected over umbilicus.
Shell chalky white with strong yellowish brown
periostracum.

Dimensions of shell : Height 6.5 mm, max.
diameter 14.1 mm: aperture height 4.8 mm.
breadth 7.0 mm.

Material examined. Eastern Atlantic. “ Por¬
cupine " 1870, stn 17a, W of Portugal. 39°39'N,
09°39'W. 1347 m: holotype (bmnh 1885.11.5.2589).

"Discovery", stn 10141. 24°34'N. 19‘41'W. 3460-
3470 m: 1 spm and 1 larva (bmnh).

Seamount, stn CP30. Gorringe Bank, 36°44'N,
11°23'W, I940-2075m: 1 young sh (mnhn).

Biacores, stn 126, Azores, 39’19'N, 33°47'W.
3360m: 1 larva (mnhn).

Biogas, stn DS 62, 47°33'N, 08°40'W. 2175m: 1
larva (mnhn).

Mozambique Channcl. Benthedi, stn 87, SE Glorieu¬
ses Islands, 11°44'S, 47°35'E, 3716 m: 2 shs (mnhn).

South Africa. “ Galathea ”, stn 192, off Durban,
32°00'S, 32*4 l'E, 3430 m: I sh (zmc).

Distribution. Eastern Atlantic Océan and
SW Indian Océan, between 40°N and 32°S, in
1940-3716m (abyssal).

Desc ription (of a shell from Mozambique
channel). Shell large, solid. depressed, consis-
ting of a globular larval shell with 3 whorls. and
1.75 planispirally coiled teleoconch whorls. Even
on best preserved benthic specimen protoconch I
and initial part of protoconch II corroded.
Protoconch II with one strong, raised spiral
thread well above suture, a second similar one,
hidden by subséquent whorl, apparent only on
terminal part of protoconch. In addition to
raised spirals, also weaker periostracal spiral
threads présent on body-whorl of protoconch, 4
such spirals between adapical suture and raised
keel, 4 between raised keel and abapical suture.
Protoconch/teleoconch transition very sharp. Te¬
leoconch with slrongly keeled whorls, giving shell
a lenticular appearance. Whorls convex above
and below keel. Body-whorl adhering to penul-
timate whorl just below keel, thus concealing
suture in a channel. Umbilicus broad and deep,
continuous with larval umbilicus. Sculpture

Larval shell measurements:

diameter no. whorls

Benthedi 87
Benthedi 87
"Galathea" 192
"Discovery" 10141
Seamount CP 30
Unmetamorphosed larvac:
holotype
Biacores 126
Biogas DS 62
"Discovery" 10141

(pm) (prot.II)

1800 ?

1920 ?

2300 2.3

1500 ?

2100 2.6

2080 2.2

2110 2.4

1810 2.5

1750 2.5

Remarks. Quinn (1983: 744) recognized
that the holotype of Seguenzia tricarinata is a
larval shell. Only a single species of Halocerati-
dae has a protoconch of this size (see measure¬
ments above and p. 159), which enables
identification of S. tricarinata. This larval shell is
superficially similar to certain nassarid larval
shells and was erroneously regarded as such by
Quinn (1983: 744). Nassarid larval shells hâve a
broad and shallow sinusigera notch above the
peripheral keel. The name tricarinata was based
on the three strong keels présent on the larval
shell. Of these only the adapical keel remains well
visible on the apex of the postlarva, the médian
keel is usually hidden just above or just below the
suture, and the basal (abapical) keel is totally
covcred by the first teleoconch whorl.

Haloceras tricarinata can be identified, both as
a larva and as a benthic snail, by its large
protoconch. If this is destroyed, the lens-shaped.
strongly keeled teleoconch, resembling certain
land snails of the genus Iherus is characteristic.
Among other species of Haloceratidae. it is only
the larval shell of H. cingulata which approaches

Source : MNHN, Paris

150

ANDF.RS WARÉN & PHILIPPE BOUCHET

Figs. 83-88. Gênera IIaimeras and Zygoceras. 83, Haloceras exquisila. hololypc. height 1.8mm. diameler 1.75mm.
84-85, Zvgoceras tropidophora, holotype, heighi 8.7mm. diameler 11.9mm. 86-87, Zygoceras sp., Monaco stn 683

heighl 1.25mm, diameler 1.7mm. 88, H. acrocomata , hololypc. heighl 2.07mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAF.

151

the size of that of H. tricarinata (range 1300-1660
vs. 1500-2300 pm). The spécifie name is most
unfortunate in view of the general appearance of
the adult shell.

The specimen from off NW Africa (“ Disco-
very", stn 10141) is still more lenticular (height
3.6 mm. diameter 10.1 mm; aperture height 3.0
mm. breadth 6.2 mm), with a weaker spiral
sculpture, and the diameter of the larval shell is
significantly smaller than in the 3 lndian Océan
snails; it may represent another species, although
the large sized larvae from the SW lndian Océan
and the NE Atlantic appear to be conspecific.

Haloceras sp. 2
not figured

Material bxamined. Australia. " Kimbla " 1984,
stn 17, NE of Sandy Cape, Queensland, 1330-1380m.

I sh (ams Cl46279).

Description. Shell small, solid, lenticular,
consisting of about 3 protoconch and slightly
more than one teleoconch whorl. Larval shell
globular, multispiral. Apical part partly corro-
ded, only sutures left, so size and sculpture of
protoconch I cannot be determined. Protoconch

II smooth, except for two spiral keels. Teleo¬
conch whorl depressed, with one sharp keel at
periphery; parts of whorl above and below keel
regularly convex. Body whorl is adhering to
penultimate whorl just below keel, giving suture
very deep appearance. Sculpture consisting of
strong spiral cords and much finer, rather indis¬
tinct incrémental fines. About 20 spiral cords
above keel, same number on base, extending into
umbilicus. Umbilicus very broad, not encircled
by a cord. Aperture ovoid, much broader than
high, angular at periphery. Outer lip sharp, thin,
inner lip slightly reflected over umbilicus.

Dimensions : Height 2.7 mm, breadth 5.05 mm;
aperture height 1.8 mm, breadth 2.70 mm.

Larval shell measurements:

diameter no.whorls

Oun) (prot.il)

1580 ca. 2.3

Remarks. — Haloceras sp. 2 is very similar to
H. tricarinata in having a single sharp peripheral
keel. In typical H. tricarinata however, the
whorls are increasing more rapidly in diameter:
at a finie more than 1 teleoconch whorl, their

diameter is already 7.2-8.5 mm, vs. only 5mm in
Haloceras sp. 2. Also, in H. tricarinata, the
peripheral keel is still sharper and more promi¬
nent. However. the larval shell characteristics fall
within the range of variation observed for typical
tricarinata. Only further material may solve the
identity of this single shell.

Haloceras trichotropoides sp. nov.

Figs 78-79, 111-112

Type material. Holotype and one pa-
ratype. both live taken, usnm 450423 and 859418.

Type locality. " Aeolis", stn 326, 136m,
off Sand Key, Florida.

Material examined. Western Atlantic (Florida).

"Aeolis", stn 326, off Sand Key. 136 m: holotype
(usnm 450423); 1 paratype (usnm 859418). — Stn 338.
off Sand Key, 156 m: 1 sh (usnm 450526). - Stn 344,
off Key West. 183m, 1 sh (USNM 450536). — Stn 369,
off Ajax Recf, 146-182m. 1 sh (usnm 450971). Stn
370, off Ajax Reef, 128-165m, 2 shs (usnm 450572). —
Stn unknown: 1 sh (usnm 878134).

Description. Shell small, solid, conical,
trochiform. consisting of 1.5 protoconch and
3 teleoconch whorls. Protoconch depressed. al-
most planispirally coiled. Protoconch I diameter
280 pm, with about 12 regular spiral cords.
Protoconch II with two strong spiral keels,
otherwise smooth: A third more basal keel.
hidden by first teleoconch whorl, is visible on a
juvénile with less than one postlarval whorl.
Teleoconch with regularly convex whorls, a deep
suture, and thick adhèrent periostracum. Sculp¬
ture consists of spiral cords and axial periostracal
lamellae. 5 spiral cords at periphery slightly
stronger than 5 above shoulder and 7 on base, of
which one encircles umbilicus and two are situa-
ted in umbilicus. Periostracal lamellae strongest
on adapical part of whorl, where forming scale-
like projections adhering to preceding whorl.
Umbilicus rather broad, open. Aperture roun-
ded; outer lip sharp, thin; inner lip not rellecled
over umbilicus, with shallow siphonal dépres¬
sion. Colour of protoconch brown, teleoconch
light amber brown.

Dimensions : Height 3.57 mm, breadth 3.06 mm:
aperture height 1.40 mm, breadth 1.38 mm.

Source : MNHN, Paris

152

ANDERS WARÉN & PHILIPPE BOUCHET

Larval shell measurements:

diameter

(Hm)

usnm 450572 670
usnm 450572 670
USNM 450526 700
USNM 450536 660
usnm 450971 610
usnm 438412 650
usnm 450423 650
usnm 450423 650

Remarks. H. triehotropoides has the smal-
lest protoconch of ail known haloceratids, with
no.whorls only a single, almost planispiral whorl in proto-

(prot.ll) conch II. Il resembles a little H. carinata , which

1.05 is partly sympatric wilh it in the West Atlantic,

1.05 but thaï species has less coarse teleoconch sculp-

l.O ture, and a globular protoconch consisting of

1.05 1.7-1.9 whorls in protoconch II.

1.0

1.05 Etymology. From the genus name Tricho-

I os tropis and the suffix -aides, having the form of;

i 05 meaning that it ressembles thaï genus.

Genus ZYGOCERAS nov.

Type species. — Zygoceras tropidophora sp.
nov.

Diagnosis. Haloceratidae with (in species
with planktotrophic development) light yellowish
multispiral protoconch. Protoconch I smooth
except for 4-6 thin spiral cords placed near
periphery. Protoconch II with two strong spiral
keels of which one is hidden below suture by
successive whorl. Teleoconch depressed with
slrongly angular whorls, narrow to closed umbi-
licus, and aperture with a swelling at base of
columellar pillar. Soft parts with characters of
the family.

Rhmarks. — Zygoceras differs from Haloceras
by the general shape and sculpture of the proto¬
conch. The protoconch is low spired and light
yellowish in Zygoceras, high spired and brown in
Haloceras. Protoconch I has only 4-6 thin spiral
cords on a smooth background in Zygoceras, vs.
anastomosing riblets and pustules in Haloceras.
In Zygoceras protoconch II has two spiral keels,
of which one only is left apparent above suture
by successive whorl; in Haloceras there are three
spiral keels, with two left apparent above suture.
The columellar swelling of the teleoconch ap-
pears to be absent in Haloceras, also in the two
larger species of comparable size ( cingulata,
tricarinata).

Although we hâve emphasized here the cha¬
racters of the protoconch of species with plank¬
totrophic development, we fully expect that
species with non-planktotrophic development.

and lherefore paucispiral protoconch, may occur
in both généra.

Etymology. From the Greek, zygos, a pair,
and the suffix -ceras, as in Haloceras-, to mean
that the new genus forms a pair wilh Haloceras,
and also to remind the two strong spiral keels.

Zygoceras biocalac sp. nov.

Figs 76-77, 123-124

Type material. Holotype in mnhn.

Type locality. Biocal, stn DW48

23°00'S, 167°29'E, 775 m. off S New Caledonia!

Material examined. New Calcdoniu. Bio¬
cal, stn DW 48, 23°00’S, I67°29'F., 775 m:
holotype (mnhn).

Description. Shell large, thin, solid, globu¬
lar, with low spire of 1.75 protoconch and 2.3
teleoconch whorls. Larval shell depressed dome-
shaped. Sculpture and limits of protoconch I not
visible on only shell available. Protoconch II with
approximately 1.25 whorls. diameter 1000 pm. One
spiral keel just above suture, a second one hidden
at suture. F.xcept for these keels, protoconch
smooth. Protoconch/teleoconch transition less
distinct than in olher haloceratids. Teleoconch
whorls convex; after First, regularly convex
whorl, two spiral keels, becoming stronger on
body whorl. In addition to keels, subtle, ill-

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

153

defined, spiral threads and strong, prosocline and
oblique, incrémental ribs. Very narrow umbilical
chink, almost closed by reflected inner lip. Aper-
ture wide, rather rounded despite spiral keels.
Outer lip thin, columellar pillar straight. Teleo-
conch white.

Dimensions: Height 7.0 mm, breadth 9.6 mm;
aperture height 4.5 mm, breadth 5.1 mm.

Remarks. — Zygoceras biocalae is a very
distinctive species, that is best identified by its
bicarinated teleoconch whorls of rapidly increa-
sing diameter, and by its almost closed umbili-
cus. Haloceras cingulata has a broad open um-
bilicus, and a sculpture of strong spiral cords; H.
exquisita and H. acroeomata hâve thin, strong
axial lamellae in addition to two very sharp keels
at the periphery. It resembles Z. tropidophora in
having a depressed protoconch with the keel
almost hidden by the next whorl, and in that
protoconch I lacks the complex sculpture of
anastomosing ridges and granules.

Etymology. From the cruise name Biocal,
a contraction for Biologie and CALédonie, during
which the material was collected in 1985.

Zygoceras tropidophora sp. nov.

Figs 28-30, 35, 84-85, 121-122

Type material. — Holotype in mnhn; one
paratype dissolved for anatomical investigation.

Type locality. Musorstom 6, stn 438,
20°23'S. 166°20'E, 780m, Loyalty ridge, E of
New Caledonia.

Material examined. Loyalty Islands (E of New
Caledonia). Musorstom 6, stn 438, 20°23'S, 166°20'E,
780 m: holotype, 1 paratype (mnhn).

Kermadec Islands. “ Akademik Nesmeyanov ", off
Curtis Island, 30’28'S, I78°37'W, 1000 m: 1 sh (nmnz
MF49882).

Description. Shell large, solid, depressed,
consisting of 2 protoconch and 1.75 teleoconch
whorls. Protoconch with very low spire. Proto¬
conch I, diameter 300 pm, sculptured by a few
spiral lines. Protoconch II, diameter 1400 pm,
consisting of 1.6 whorls, smooth with sharp
spiny keel at periphery, just above suture; pro-
jecting spines calcareous. not periostracal. Teleo¬
conch whorls very angular, of rapidly increasing

diameter. Body-whorl very large. 2 strong keels
at periphery give the whorl a roughly square
cross-section. Shoulder and base moderately
convex. Beside 2 keels, sculpture consists of thin
weak spiral cords and incrémental ridges; ridges
stronger at periphery and fading on shoulder and
base. Between pcriphcral keels, spiral and axial
sculpture of equal strength; intersections forming
fine sharp décussation. No umbilicus. Aperture
strongly angular, with basal part considerably
drawn out, slightly flaring. Outer lip thin, sharp.
Inner lip indistinct. Columellar pillar strong.
excavated at mid aperture height, slightly twisted
below. Colour of protoconch light yellowish
brown, teleoconch white.

Dimensions: Height 8.7 mm, breadth 11.9 mm;
aperture height 7.1 mm, breadth 6.8 mm.

Larval shell measurements:

diameter

no. whorls

(pm)

(prot.10

holotype

1400

1.6

nmnz 49882

1450

1.8

Remarks.

- Zygoceras

tropidophora can

hardly be confused with any other species of
Haloceratidae. Halocears cingulata differs by
having much more depressed whorls and aper¬
ture, weaker peripheral keels, and a broad open
umbilicus. Z. hiocalae has a similar low proto¬
conch, but with fewer whorls and without pro-
jecting spines on the spiral keel. Furthermore it
has a small but distinct umbilicus, weaker spiral
sculpture and a more rounded aperture.

The protoconch of Z. tropidophora stands out
among ail gastropod larval shells we hâve exa¬
mined in having a keel of solid calcified projec-
ting spines (not periostracal bristles as is found in
a number of Tonnoidea). In that respect it
resembles a miniature Cochliolepis gruveli from
West Africa (see Adam & Knudsen 1969: fig.33).

Etymology. — From the Greek tropis. keel,
and phoreus, carrier; to remind of the teleoconch
sculpture. Tropidophora is also a genus of land
snails from Madagascar, which has a superficial
ressemblance with Zygoceras.

Source : MNHN, Paris

154

ANDERS WARÉN

PHILIPPE BOUCHET

Figs. 89-96. Protoconchs of Haloceras. 89-90, H. exquisila, holotype. 91-92, II. rugosa, holotype. - 93 H
acrocomata, holotype. 94, H. acrocomata. AMS C 147279. 95-96, //. phaeocephala. paratype. Scale lincs 250 uni

(89, 91, 93. 95) and 50 pm (90, 92, 94, 96).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HA LOCER ATI DA E

155

FiCis. 97-104. Protoconchs of Haloceras. 97-99, II. mediocostata, Icctolypc. 100-102. H. galerila, holotype. 103,
II. Iieliptyx. holotype. 104, Haloceras sp.l. ams C 157269. Scalc Unes 250 pm (97-98. 100-101. 103-104) and 50 um
(99, 102).

Source : MNHN, Paris

156

ANDERS WARÉN

PHILIPPE BOUCHET

Figs. 105-112. — Protoconchs of Haloceras. 105-106, II. carinata, syntype of Salarie Ha ainsi rie la.
unmetamorphosed larva. incal stn DS3. - 100-110, II. spinosa. holotypc. 111-112, //. tricha
Scale lines 250 pm (105-107, 109, 111) and 50 pm (108, 110. 112).

107-108. H. carinata,
ipoiilc.s. usnm 450526.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

157

Figs. 113-120. Protoconchs of H aimeras. 113, H. cingulata, " Thalassa ” stn Z435. 114-115, H. cingulata. Seamount

sln CP30. 116-118, H. taxa. Seamount sin CP30. 119-120, H . tricarinata . “ Discoverv " sin 10141. Scale Unes 250

Hm (113-114. 116-117. 119) and 50 nm (115. 118. 120).

Source : MNHN, Paris

158

ANDERS WARÈN

PHILIPPE BOUCHET

Figs. 121-126. Proloconchs of Zygoceras. 121 - 122 , Z. iropidophora, hololypc 123 - 124 . Z. biocalae, hololvne
125 - 126 . Zygoceras sp.. Monaco sln 683. Scalc Unes 250 pm (121, 123, 125) and 50 pm (122. 124. 126).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

159

Zygoeeras sp.

Figs 86-87. 125-126

Material examiner. Kastcrn Atlantic. Monaco,
stn 683, Azores, 38°20'N, 28 < ’05'W. 1550 m: 1 young sh
(mom).

Remarks. A very characteristic juvénile
with 0.6 postlarval whorl, which cannot be

identified with any of the described species.
Protoconch I, diameter 350 pm, has a sculpture
of spiral cords on the nucléus and the abapical,
coiled part, leaving a smooth adapical zone.
Protoconch II. diameter 980 pm. is low-spired
and has 1.25 whorls with only 2 keels instead of
3 as in Halocercis. One keel is visible above the
suture, a second one concealed by the suture.
The teleoconch has two strong keels at periphery.

IDENTIFICATION: SUMMARY OF LARVAL SHELL CHARACTER1STICS

The multispiral protoconch of the Halocerati¬

Table 2.

- Larval shell characteristics

dae offers very good characters for identification

of Haloceratidae

of the species. To facilitate comparisons, we hâve

compiled the characteristics of ail species below.

diameter(gm)

no.whorls

arranged from the smallest to the largest larval

prot.ll

shell.

range

mcan

range r

trkhotropoides

610-700

657

1.0-1.05

1.04

8

spinosa

675-720

695

1.2-1.3

1.27

3

heliptyx

770

770

?

1

carinata

700-920

817

1.7-1.9

1.82

16

exquisita

870

870

2.1

2.1

1

phaeaccphala

890-1060

967

1.8-2.0

1.87

6

acrocomata

940-1000

970

1.4-1.6

1.49

4

bioealae

1000

1000

1.25

1.25

1

galerita

1080

1080

2.1

2.1

1

taxa

980-1200

1118

1.7-1.8

1.76

8

rugosa

1100

1100

2.1

2.1

I

mediocostata

1200-1300

1240

2.0-2.1

2.06

5

Iropidophora

1400-1450

1425

1.6-1.8

1.7

2

cingulata

1300-1660

1466

2.1-2.3

2.19

7

tricarinata

1500-2300

1930

2.2-2.6

2.42

9

Not cnlcrcd in table: japonica, millestriata.

ACKNOWLEDGEMENTS

We thank the curators of the institutions
which hâve loaned us material of Haloceratidae
over the last 10 years: W. Ponder and I. Loch
(ams). B. Marshall (nmnz), J. McLean (lacm).
R. Houbrick and the late J. Rosewater (usnm),
E. Gittenberger (rmnh). T. Okutani (Tokyo)
and K. Way (bmnh). We also thank the leaders
and participants to the various deep-sea expédi¬
tions in New Caledonia that collecled halocera-

tids: C. Lévi, C. Monniot, B. Métivier and P.
Lozouet (mnhn). and B. Richer de Forges
(orstom, Nouméa). A. Warén also wants to
thank ams for a position as visiting research
scientist during which much of this material was
found.

V. Héros (mnhn) and M. Segonzac (centob,
Brest) sorted much of this material. S. Gofas
prepared figure 64.

Source : MNHN, Paris

160

ANDERS WARÉN & PHILIPPE BOUCHET

We also want lo thank A. Graham, G.
Haszprunar and W.F. Ponder who read and
commented this manuscript.

For technical assistance in the préparation of

this work, we thank C. Hammar and A. Heds-
trôm (smnh) and D. Guillaumin (Centre de
Microscopie, cnrs, Paris).

REFERENCES

Adam, W. & Knudsen, J.. 1969, Quelques genres
de Mollusques Prosobranches marins inconnus ou
peu connus de l'Afrique occidentale. Bull. Inst. r.
Sci. nal. Belg.. 44(27): 1-69.

Boss, K. J., 1982. Mollusca. In: S. P. Parker (ed.).
Synopsis and Classification of Living Organisms.
McGraw Hill Book Co., New York. Vol. 1: 945-
1166, Vol. 2: 1092-1096.

Bouchet. P., 1976. Mise en évidence d'une migra¬
tion de larves véligères entre l’étage abyssal et la
surface. C. r. Itehd. Séanc. Acad. Sci.. Paris. (D) 283:
821-824.

Bouvier, E.-L., 1887. — Système nerveux. Morpho¬
logie générale et classification des gastéropodes
prosobranches. Annls. Sci. nal. Zoo!., (7)3:1-510.

Dall, W.H.. 1889. Reports on the resulls of
dredging, under the supervision of Alexander Agas-
siz. in the Gulf of Mexico (1877-78) and in the
Caribbean Sea (1879-80), by the US coast survey
steamer « Blake », Lieut.-Commander C.D. Sigsbee,
U.S.N. and Commander J.R. Bartlett, u.s.N.
commanding. XXIX. -Report on the Mollusca. Part
II. Gastropoda and Scaphopoda. Bull. Mus. camp.
Zool. , 18: 1-491.

Dall, W.H., 1927. Small shells from dredgings off
the southeast coast of the United States by the
United States Fisheries Steamer « Albalross », in
1885 and 1886. Proc. U.S. nain. Mus.. 70(18): 1-134.

Dautzenberg, P. & Fischer, H., 1896. Dragages
effectués par P « Hirondelle » et par la « Princesse-
Alice»: 1. Mollusques Gastéropodes. Mém. Soc.
zool. Fr.. 9: 395-498.

Dautzenberg, P. & Fischer, H., 1897. Dragages
effectués par 1’ « Hirondelle » et par la « Princesse-
Alice» 1888-1896. Mém. Soc. zool. Fr.. 10: 139-234.

Dautzenberg. P., 1927. — Mollusques provenant des
campagnes scientifiques du Prince Albert Ier de
Monaco dans l’Océan Atlantique et dans le Golfe de
Gascogne. Resuit. Camp, scient. Prince Albert I. 72:
1-407.

Dell. R. K. & Ponder, W.F., 1964. Sonie notes on
the gastropod Zelippistes benhami (Suter), Rec.
Dom. Mus. Wellington. 5: 63-72.

Giese, M., 1915. Der Genitalapparat von Calyp-
traea sinensis Lin.. Crepidula unguiformis Lam. und
Capulus hungaricus Lam. Z. wiss. Zool.. 114:169-
231.

Graham, A.. 1954. — The anatomy of the proso-
branch Trichotropis horealis Broderip & Sowerby,
and the systematic position of the Capulidae J
mar. biol. Assoc. U.K.. 33: 129-144.

Haller, B.. 1893. Die Morphologie der Prosobran-
chier. IV. Morph. Jh., 19:553-591.

Houbric k, J.R. & Fretter, V., 1969. Some aspects
of the functiona! anatomy and biology of Cymatium
and Bursa. Proc, malac. Soc. London. 38:415-429.

Jeffreys, J.G., 1883. On the Mollusca procured
during the « Lightning » and « Porcupine » expédi¬
tions 1868-70. Part VI. Proc. zool. Soc. London
1882: 88-149.

Jeffreys, J.G., 1885. On the Mollusca procured
during the « Lightning » and « Porcupine » expédi¬
tions 1868-1870. Part IX. Proc. zool. Soc. London
1885: 27-63.

Lacaze-Duthiers. H. de, 1872. Otocystes des
mollusques suite, troisième type. Archs Zool exn
gén., (3)9:43-79. y '

Lacaze-Duthiers, H. de, 1901. Le système nerveux
du Cabochon Capulus hungaricus. Archs Zool exn
gén.. (3)9:43-79. ' ' '

Morton. J.E., 1952. The rôle of the crystalline
style. Proc, malac. Soc. London. 29: 85-92.

Okutani, T., 1964. - Report on the archibenthal and
abyssal gastropod mollusca collected from Sagami
Bay and adjacent waters by the R.V. Soyo-Maru
during the years 1955-1963. ./. Fac. Sci Tokvo
Unix.. . (2) 15: 371-447.

PILKINGTON, M.C., 1974. The eggs and hatching
stages of some New Zcaland prosobranch mollusrs
J! R. Soc. N.Z.. 4:411-431.

PILKINGTON, M.C., 1976. Descriptions of veliger
larvae of monotocardian gastropods occurring in
Otago plankton hauls. J. molluscan Stud.. 42:337-

Ponder, W., 1985. A review of the généra of
Rissoidae (Mollusca: Mesogastropoda: Rissoacea)
Rec. Auslr. Mus., suppl. 4: 1-221.

Ponder, W. & Waren, A., 1988. Classification of
the Caenogastropoda and Hetcrostropha- A list of
the family-group naines and higher taxa. Malac
Review, suppl. 4: 288-326.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NEW FAMILY HALOCERATIDAE

161

Quinn, J. F., 1979. Biological results of the Univer-
sity of Miami deep-sea expéditions. 130. The syste-
matics and zoogeography of the gastropod family
Trochidae collected in the straits of Florida and its
approaches. Malacologie !, 19: 1-62.

Quinn, J.F., 1983. A révision of the Seguenziacea
Verrill, 1884. I. Summary and évaluation of the
superfamily. Proc. Biol. Soc. Wash.. 96(4): 725-757.

Thiele, J., 1929-34. — Flandbuch der systematischen
Weichtierkunde. I. G. Fischer, Stuttgart. 1023 pp.

Thorson. G., 1965. A ncotenous dwarf-form of
Capulus ungaricus (L.) (Gastropoda, Prosobranchia)
commensalistic on Turrilella commuais (Risso).
O plie lia, 2:175-210.

Verrill, A.E.. 1884. Second catalogue of Mollusca
recently added to the fauna of the New England
coast and the adjacent parts of the Atlantic, consis-
ting mainly of deep-sea species with notes on others
previously recorded. Trans. Corn. Acad. Sci. Arts, 6:
139-294.

Warén, A., 1979. Marine Mollusca described by
John Gwyn Jeffreys, with the location of the type

material. Conchological Society of Great Britain
and Ireland, Spécial Publication 1: 1-60, 8 plates.

Warén, A., 1984. An anatomical comparison of
Eulima and Pvramidelloides with a révision of the
species of Pyramidelloides (Mollusca, Prosobran¬
chia, Eulimidae). Zoologica Scripta. 12:273-294.

Warén, A. & Bouchet, P., 1989. Laubierinidae
and Pisanianurinae (Ranellidae). two new deep sea
taxa of the Tonnoidea (Gastropoda: Prosobran¬
chia). Veliger. 33: 56-102.

Warén, A. & Ponder. W. F., (in press). — New
species, anatomy and systematic position of the
hydrothermal vent and hydrocarbon seep family
Provannidae n.fam. (Cacnogastropoda). Zoologica
Scripta

Wenz, W., 1938-44. Gastropoda. I. Allgemciner teil
und Prosobranchia. Handbuch der Palâozoologie,
6:1-1639.

Yonge, C M., 1962. On the biology of the meso-
gastropod Trichotropis cancellala Hinds. a benthic
indicator species. Biol. Bull., 122: 160-181.

Source : MNHN, Paris

Source : MNHN, Paris

ULTATS DES CAMPAGNES MUSORSTOM, VOLUME 7

RÉSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSUL

5

Mollusca Gastropoda :

Four new rissoinine species (Rissoininae)
from deep water in the New Caledonian région

Willy J. SLEURS*

Koninklijk Bclgisch Instituut voor Natuurwetcnschappen
Recent Invertebrates Section
Vautierstraal, 29
B 1040 Brussels
Belgium

ABSTRACT

Four new spccies, belonging to the subfamily Rissoininae
(Ncotacnioglossa : Truncatelloidca : Rissoidae), are dcscri-
bed from deep water in the New Caledonian région : Rissoina

(Rissoina) bouchai sp. nov., R (R.) longispira sp. nov.,
Zebina (Zebina) réclina sp. nov. and Z. (Z.) refusa sp. nov.
An analomical description of R. bouchai is given.

RÉSUMÉ

Mollusca Gastropoda : Quatre espèces nouvelles de Rissoi-
ninac (Rissoidae) des eaux profondes néo-calédoniennes.

Quatre nouvelles espèces de la sous-famille Rissoininae
(Neotacnioglossa : Truncatelloidea : Rissoidae) sont décrites :
Rissoina (Rissoina) bouchai sp. nov., R. (R.) longispira sp.

nov., Zebina (Zebina) réclina sp. nov. et Z. (Z.) refusa sp.

Rissoina est présent jusqu'à 700 m de profondeur, ce qui
représente les occurences les plus profondes actuellement
connues.

* The author is granted by the “ National Fund for Scicntific Research (Belgium) ".

Sueurs, W. J.. 1991. Mollusca Gastropoda : Four new rissoinine species (Rissoininae) from deep water in the New Caledonian région.
In A. Ckosnikk & P. Bouchet (cds). Résultats des Campagnes Musorstom, Volume 7. Mém. Mus. nain. Hisl. nai., (A). 150 : 163-178. Paris
ISBN : 2-85653-180-6.

Source : MNHN, Paris

164

WILLY J. SLEURS

INTRODUCTION

Most of the material reported here was collected
during the Biocal Expédition, conducted in 1985
aboard the R. V. “ Jean-Charcot ", undcr the
direction of Prof. C. Levi. Specimens were sorted
on board by P. Bouchet, B. Metivier and B.
Riciier de Forges, and residues were saved for
further sorting at Centob, Brest, under the
supervision of M. Skgonzac. For further infor¬
mation on the expédition, see Richer de Forges
(1990).

Additional material collected during the Smib 3
Expédition (1986), south of Ile des Pins and on
the guyots of the Norfolk ridge is included.
Furthermore also a small collection, made aboard
the “ Kimbla " in 1971 and housed in the ams, in
included.

This paper represents the first study on the

deep water Rissoininae from New Caledonia : ail
previous studies on New Caledonian rissoinines
refer to shallow water species. This study forms
part of a species review on the Rissoininae,
currcntly in progress.

Abbreviations of institutions :
ams : Australian Muséum, Sydney.
bmnh : The Natural History Muséum, London.
kbin : Koninklijk Belgisch Instituut voor
Natuurwetenschappen, Brussels.

I.ACM : Los Angeles County Muséum.
mnhn : Muséum national d'Histoire naturelle,
Paris.

nmnz : National Muséum of New Zealand,
Wellington.

SYSTEMATIC ACCOUNT

Superorder CAENOGASTROPODA Cox, 1959
Order NEOTAENIOGLOSSA Haller, 1882
Superfamily TRUNCATELLOIDEA Gray. 1840
Family RISSOIDAE Gray, 1847
Subfamily RISSOININAE Stimpson, 1865

The généra of the family Rissoidae are revised sed in that paper is followed here ; therefore we
by Pondhr (1985), and the classification propo- refer to that paper for generic diagnoses.

Genus RISSOINA

Rissoina (Rissoina) boucheli sp. nov.

Figs 1-2, 3 a-c, 5-11

Type material. Ail from Biocal, stn DW 44,
30.VI1I.1985. Holotype : empty shell in mnhn.
Paralypes ; 30 specimens (some of them with
dried animal) in mnhn. One paratype in the

d’Orbigny, 1840

following institutions or muséums : ams, kbin,
lacm and nmnz.

Type locality. New Caledonia, 22"47' S
167°14' E, 440-450 m.

Material examined (Apart from type material).
New Caledonia. Biocae : stn DW 33, 23" 10' S. 167“ 10' E

Source : MNHN, Paris

MOLLUSCA GASTROPODA : DEEP-WATER R/SSOINA

165

Fig. 1. Rissoina (s. s.) boucheli sp. nov. : a-c : shclls and f-h : protoconchs. a holotype. New Caledonia. Biocal stn DW
44 (mnhn) : b soulh New Caledonia. Smib 3 stn DW 22 (mniin) : c - paratype. New Caledonia. Biocal stn DW 44
(mnhn) ; d New Caledonia. Biocal stn DW 77 (mnhn) ; e south New Caledonia. Smib 3 stn DW 22 (mnhn) : f
paratype. New Caledonia. Biocal stn DW 44 (mnhn) : g paratype. New Caledonia. Smib 3 stn DW 22 (samc
spécimen of Fig. I e) ; h New Caledonia. Biocal stn 77 (mnhn).

Scale : a-e = 2 mm ; f-h = 0.5 mm.

Source : MNHN , Paris

166

W1LLY J. SLEURS

675-680 m, 29.V111.1985 : 6 spcc. (mnhn). Stn DW 41,
22“45'S. 167° 12' E. 380-410m. 30.VIII.1985 : 1 spec.
(mnhn). Stn DW 43, 22°46'S, 167°15'E, 400 m.
30.V1I1.1985 : 4 spec. (mnhn). — Stn DW 44, 22°47' S.
167° 14' E, 440-450 m, 30.V111.1985 : 90 spec. (with
animais preserved in alcohol) (mnhn). Stn DW 46,
22"53'S. 167“ 17' E, 570-610m, 30.VII1.1985 : 2 spec.
(mnhn). Stn DW 56, 23°35'S, 167°12'E. 705-
695 m. 01.IX. 1985 : 3 spec. Stn DW 77, 22“15' S,
167° 15' E, 440 m, 05.1X. 1985 : 75 spec. (mnhn).

Smib 3, R. V. “ Vauban " : stn DW 10, 235 m,
21.V.1987 : 1. spec. — Stn DW 21, 22°59' S. 167°19' E,
525 m, 24.V.1987 : 1 spec. (mnhn). — Stn DW 22,
23°03'S, 167° 19' E, 503 m. 24.V.I987 : 73 spec. (mnhn).

Hmas " Kimbla ", stn K 4-71-3, 22°50' S, 167°34' E
(approx. 7 km S. of I. des Pins), 275 m, coral sand
bottom, 08.V. 1971, coll. P. H. Colman & J. Paxton :
3 spec. (ams, cl 53935) ; Ibidem. 274 m : 3 spec. (ams,
cl 53936).

Description. Shell (Fig. 1 a-e) : moderately
large (length 9.9 mm in holotype), solid, more or
less elongate cortical ; last whorl moderately
angulate to subangulate at the periphery.

Protoconch (Fig. 1 f-h) : of non-planktotrophic
larval type (and probably with intracapsular
metamorphosis), moderately elongate conical to
subcylindrical, glossy, of about 2 smooth whorls ;
transition to teleoconch abruptly with a straight,
non-thickened margin.

Teleconch : of about 7 to 8 whorls ; spire
whorls almost fiat, weakly to moderately angu¬
late just below and/or above the deeply to
moderately impressed, weakly undulating sutures.

Axial sculpture of slightly opisthocline axial
ribs, the latter variable in strength, ranging from
very prominent, sharp, distantly spaced, to weak,
densely spaced and rounded ; axial sculpture
usually somewhat more prominent on spire
whorls than on last whorl, but sometimes equal
in strength throughout ; axial ribs on last whorl
somewhat weaker below periphery, but conti-
nuous to peristome.

Spiral sculpture very variable, ranging from
microscopie, irregular and irregularly spaced
scratches (Fig. 3 a-c), to moderately prominent,
more or less regularly and densely spaced spiral
threads or weak spiral ribs ; spiral sculpture,
when présent, mostly more prominent on penul-
timate and last whorl than on the early spire
whorls.

Aperture : moderately large, D-shaped to
lenticular or auriculiform ; columellar side mode¬
rately concave ; anterior channel narrow, short.

shallow ; outer lip with a prominent interior
swelling near the transition to the anterior
channel ; posterior channel very short, triangu-
lar ; outer lip slightly opisthocline in profile with
a moderately wide, rounded, prominent varix
externally.

Colour : yellowish white with large orange
spots on last and penultimate whorl or with a
rather wide orange spiral band below the suture ;
some specimens white throughout.

Operculum (Fig. 2 e-f) : thick, with weakly
curved, hollow peg ; muscle-attachment area
sausage-shaped.

Radula (Fig. 2 a-d) : central teeth of taeniglos-
satte radula with the formula (Ponder, 1985 : 10) :
3-4 + I + 3-4 , . . ,

----- t ; ventral margin with strongly-

developed U-shaped extension ; latéral margins
making an angle of ca. 30° with dorsoventral
axis. Latéral teeth 7-9 + 1 + 3-4. Inner marginal
teeth with 7-11 sharp cusps on distal half of
outer edge and with some weak cusps just below.
Outer marginals with weak cusps on outer and
inner edge.

Head-foot characlers : cephalic tentacles usually
covered by a thin sheath (probably an artifact
due to préservation), the latter with deeply
indented margins ; right palliai tentacle narrow,
rather long, simple ; left palliai tentacle deeply
bifurcate, consisting of 2, rather wide, lobes ;
metapodial tentacle not observed in preserved
specimens.

Mande cavity (Fig. 11) : mean ctenidium
length : 2.45 mm (n = 5) ; mean number of gill
filaments : 45 (n = 5) ; gill filaments rather
short, longest gill filament measuring approx.
0.25 mm ; osphradium somewhat shorter than
ctenidium, consisting of a rather thick, wide
undulating (probably due to contraction) main
ridge ; latéral ridges very narrow, hidden beneath
the mean ridge ; hypobranchial gland inconspi-
cuous.

Digestive System (Figs 5, 7, 11) : mouth
opening between two fleshy lips into a rather
long buccal tube ; one pair of jaws in anterior
third of buccal tube ; salivary glands simple
single tubes, not reaching the nerve ring and with
distal part folded ; oesophagus rather uniform in
structure from its departure from buccal cavity
to the opening into the anterior chamber of the
stomach ; oesophagus with about 10 folds inter-
nally. Stomach occupying about one whorl ;

Source : MNHN, Paris

MOLLUSCA GASTROPODA : DEEP-WATER RISSOINÂ

167

Fig. 2. ^ Sem micrographs of radula and opcrculum of Rissoina (s. s.) bouchai sp. nov., (kbin stub 77 E-F). New Calcdonia.
Bkk al stn DW 44. a centrais, laterals and inner marginals ; b laterals ; c central ; d distal part of outcr
marginal ; e inner sidc of opcrculum ; f latéral aspect of inner side of operculum.

Scalc : a, b. c = 0.02 mm; d = 0.01 mm ; e. f = I mm.

Source : MNHN , Paris

168

WILLY J. SLEURS

Fig. 3 a-c. — Variation in microsculpture of teleoconch (penullimate whorl) of Rissoina <Rissoina) bouchai sp. nov. ■ a

paratypc. New Calcdonia. Biocai. stn DW 44 (mnhn) : b New Caledonia, Biocal stn DW 77 (mniin) ; c New
Caledonia, Biocai. stn DW 22 (mnhn).

Fig. 3 d-f. Rissoina (Rissoina) longispira sp. nov. : d holotypc (mnhn); e paratypc (immature spécimen), New
Caledonia. Biocal stn DW 33 (mnhn); f protoconch (samc spécimen as Fig. 3c).

Scale : a. b. c = 0.1 mm ; d, e = 2 mm ; f = I mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : DEEP-WATER RISSOINA

169

FlCi. 4a-d. Zebina (Zebina) relusa sp. nov. : a. b
Calcdonia. Biocal sln DW 38 (mnhn).

Fig. 4 e-h. Zebina (Zebina) réclina sp. nov. : e, f
Calcdonia. Biocal stn DW 44 (mnhn).

Scalc a, b = 1 mm ; c, e, f, g = 2 mm ; d. h

holotype (mnhn) ; c, d
holotype (mnhn) ; g, h
= 0.1 mm.

Shell and protoconch of paratypc. New
shcll and protoconch of paratype. New

Source : MNHN, Paris

170

WILLY J SLF.URS

Fig. 5. Anatomy of uncoiled female of Rissoina (s. s.) boucheli sp. nov., New Calcdonia, Biocal sln DW 44.

(Abbreviations : ac anterior chambcr of stomach ; bc bursa copulatrix ; cm columellar muscle ; et -

cephalic tentacle ; dg digestive gland ; e cyc ; cme end of mantle cavity ; fp faecal pellct ; int intestine ; log

— lower oviduct gland ; oes oesophagus ; ov — oviduct ; ovy ovary ; pc posterior chambcr of stomach ; rpt -
right palliai tentacle ; ss style sac ; uog — upper oviduct gland).

stomach morphology typical of genus : stomach/
style sac ratio (as defined by Ponder, 1984 :
13) : 13 (n = 3) (due to the very elongate
posterior chamber) ; length/width ratio (as defi-
ned by Ponder, 1985 : 13) : 3.7; stomach
containing foraminiferan material and fragments
of filamentous algac ; style sac short (crystalline
style not observed) ; gastric shield broadly trian-
gular ; digestive gland dark brown, occupying
about 2 1/2 whorls, not anterior to anterior
chamber of stomach ; intestine very thin-walled,
very wide in mid-section, filled with numerous
cylindrical fecal pellets with rounded anterior
and posterior ends.

Female reproductive System (Figs 5-6) : ovary
very short ; oviduct thin-walled ; upper oviduct
gland large, more or less bilobcd, subequal to
length of lower oviduct gland, oval in section,
with a slit-like lumen ; bursa copulatrix rather
small, between lower and upper oviduct gland,
partly overlying both glands at the right side ;
séminal réceptacle not observed (probably over-
looked or inconspicuous when not filled with
sperm) ; sperm duct strongly muscular. narrow,
lying along left side of lower oviduct gland, only
very weakly expanded near anterior end. One
mature female was found with normal female
reproductive organs and with a non-functional

Source : MNHN, Paris

MOLLUSCA GASTROPODA : DEEP-WATER RISSOINA

171

pénis, the latter very short as in immature male
specimens.

Male reproductive System (Figs 7-8) : testis
occupying about 1 whorl, overlying the digestive
gland ; séminal vesicle highly coiled, strongly
expanded, with posterior half folded over poste-
rior half of posterior chamber of stomach ;
anterior half covered by digestive gland ; viscéral
vas deferens. passing along stomach ; prostate
gland open, elongate, thin. Pénis occupying most
of mantle cavity, large with a wide, spoon-shaped,
distal lobe ; penial groove open, the latter ter-
minating as a somewhat expanded gutter just
above the spoon-shaped lobe ; one margin of the
latter weakly denticulate ; ventral side of spoon-
shaped lobe almost fiat sided ; dorsal side of lobe
with a médian crest, the latter with a denticulate
summit.

Central nervous system (Figs 9-10) : cérébral
ganglia joined by a rather long commissure for
genus ; RPG ratio (as defined by Davis et al.,
1976 : 263) : 0.6 ; pleural ganglia separated from
cérébral ganglion by a rather deep constriction ;

statocysts on the posterior edge of the pedal
ganglia.

Shell dimensions and sculpture counis : See
table 1.

Variation. — Rissoina boucheti sp. nov. shows
considérable interpopulation variation with res¬
pect to shell sculpture, but is strikingly uniform
within the same population. In the type sériés
(Fig. 1 a, c), the axial ribs are very prominent,
regularly spaced and sharp, while the spiral
sculpture is almost absent except for some
irregularly spaced scratches ; in specimens of
other populations the axial sculpture ranges
from moderately prominent (Fig. 1 d) to very
weak, rounded, closely spaced ribs (Fig. 1 b, e) ;
the spiral sculpture ranges from irregularly spaced
microscopie scratches (Fig. 3 a) to more or less
regularly spaced, moderately prominent spiral
threads or fine spiral ribs (Figs 1 d ; 3 b, e).

In the type sériés the whorls are moderately
angulate below the suture (Fig. la, c) ; in other

Fig. 6. - Fcmalc genilalia of Rissoina (s. s.) boucheti sp. nov. (viscéral oviduct and ovary omitted). New Caledonia. Biocal
stn DW 44.

(Abbreviations : bc bursa copulatrix ; gp génital porus ; log — lower oviduct gland ; ov — oviduct : sd —
sperm duel ; vc ventral channel ; uog — upper oviduct gland).

Source : MNHN , Paris

172

WILLY J. SLEURS

PC

2 mm

Fig. 7. — Anatomy of an uncoilcd male of Rissoina (s. s.) boucheli sp. nov.. New Calcdonia, Biocal stn DW 44. (digestive
gland partly removed to show the position of the séminal vesicle).

(Abbreviations : ac anterior chambcr of stomach ; cm columellar muscle ; et — cephalic tcntacle ; dg -
digestive gland ; c eye ; emc end of mantle cavity : fp faecal pcllct ; int intestine ; oes ocsophagus ; p
pénis ; pc posterior chambcr of stomach ; pr prostate gland ; pvd palliai vas deferens ; rpt right palliai

tentacle ; ss - style sac ; sv - séminal vesicle ; t testis ; vvd viscéral vas diferens : part of viscéral vas déferons

indicated by dotted lines was never found, but it is here marked in ils presumed position).

populations we found specimens with no or only
very weakly angulate whorls (Fig. 1 b, d, e).

Etymology. — This species is named after Dr

Philippe Bouchet of the mnhn, who made this
New Caledonian collection of rissoinines avai-
lable for examination, and who was one of the
collectors of this new species.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : DEEP-WATER RISSOINA

173

Fig. 8. Pénis of Rissoina (s. s.) boucheli sp. nov., New Caledonia. Biocal sln DW 44 (a dorsal aspect. " in situ " ;
b ventro-lateral aspect after stretching ; c ventral aspect of spoon-shaped distal lobe : d transverse sections a. b.
C and d. taken at positions indicated in Fig. 4 a : penial gland dark).

(Abbreviations : g gutter ; pgl penial gland ; pgr — penial groove).

Discussion. Rissoina (s.s.) boucheti sp.
nov. is very similar to Rissoina aupouria Powell,
1937, from Three Kings Islands (New Zealand),
but diflers in the protoconch being more conical,
in having a more angulate last whorl and in
having a prominent swelling near the base of the
inner side of the outer lip ; furthermore the spiral
sculpture is much weaker in R. aupouria and
consists of numerous, very closely spaced, micros¬
copie striations.

The new species is also very similar to three

species, formerly included in the genus Siiva
Hedley, 1904, from south-eastern Australia. Pon-
der (1985 : 82), however, considered the species
of Siiva do not difTer sufficiently from Rissoina
(s.s.) to recognize Stiva even as a distinct
subgenus of Rissoina.

R. (R.) boucheli difTers primarily from R. (R.)
J'erruginea (Hedley, 1904), R. (R.) royana (Ire-
dale, 1924) and R. (R.) nielseni (Laseron, 1950)
in having a prominent ridge or swelling on the
inner side of the outer lip, near the transition to

Source : MNHN, Paris

174

W1LLY J. SLEURS

Fig. 9. — Nervous System (dorsal aspect) of Rissoina (s. s.)
boucheti sp. nov., New Caledonia, Biocal stn DW 44.

(Abbreviations : bg — buccal ganglion ; cm — columel-
lar muscle; leg left cérébral ganglion; lplg left
pleural ganglion; oes — oesophagus; plspo — pleural
supraoesophageal connective ; r — radula ; reg right
cérébral ganglion ; rplg — right pleural ganglion ; sbcg —
suboesophageal ganglion ; spog — supraoesophageal gan¬
glion).

Table 1. — Rissoina (Rissoina) boucheti sp. nov. Shell
dimensions and sculpture counts (i. : shcll length ; ls :
length of spire ; D : shcll diameter ; Dp : diameter of last
whorl of protoconch ; Nax : number of axial ribs on last
whorl ; Naxp : number of axial ribs on pcnultimate whorl ;
x : mean ; SD : standard déviation ; o.r. : observed range ;
n : number of specimens).

L Ls D
(mm) (mm) (mm)

Nax

Naxp

Dp

No. whorls

Holotype
(Fig. 1 a)

9.9 6.8

3.6

24

21

0.73

7

Paratypcs (in M

nhn except where cxplicitcly stated)

10.4 7.3

3.8

22

0.75

7

NMNZ

10.0 6.7

3.9

21

21

?

9.8 6.8

3.5

21

18

0.70

7

AMS

9.7 6.6

3.7

21

18

0.75

7

9.6 6.8

3.5

27

26

0.73

7 1/4

9.6 6.5

3.7

25

16

0.75

7

9.4 6.5

3.5

24

20

0.75

7

9.4 6.3

3.7

22

20

0.75

6 3/4

7

9.2 6.3

3.6

30

19

0.75

Stn DW 44 (n

= 32)

x

9.65 6.61

3.64

23.9

19

7.0

SD

0.33 0.28

0.17

3.0

2.3

0.2

O.R.

9.1- 6.2-

3.0-

19-

16-

6 1/2-

10.4 7.3

3.9

31

26

7 1/2

Stn DW 22 (n

= 22)

x

11.2 8.0

3.8

32.2

26.8

8.1

SD

0.47 0.35

0.15

4.1

4.4

0.3

10.8- 7.6-

3.5-

25-

19-

11.8 8.5

4.1-

41

40

8 3/4-

Stn DW 77 (n

= 22)

x

8.8 5.8

3.2

23

19

7.3

SD

0.36 0.9

0.1

1.9

1.5

0.3

OR

8.1- 5.5-

3.1-

20-

17-

9.2 6.6

3.4-

26

21

the anterior channel. R. (R.) boucheti diflfers
anatomically from R. (R.) ferruginea in having
an osphradium consisting of a thick undulating
ridge, while the osphradium in the latter is of the
bipectinate type ; futhermore R. (R.) ferruginea
has a simple, very short left palliai tentacle, while
the latter is bifurcate and well developed in R.
(R.) boucheti sp. nov.

Fig. 10. — Nervous System (latéral aspect) of Rissoina (s. s.)
boucheti sp. nov.. New Caledonia, Biocal stn DW 44
(buccal ganglia omitted and left pleural ganglion covered
by left cérébral ganglion).

(Abbreviations : btn — bulbus of tentacular nerve ; leg

— left cérébral ganglion; Ipg — left pedal ganglion;
plspoc plcuro-supraocsophagcal connective ; reg —
right cérébral ganglion ; rpg — right pedal ganglion ; rplg

— right pleural ganglion; spog supraoesophageal

ganglion) ; stc — statocyst ; stl — statholith ; tn _

tentacular nerve).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : DEEP-WATER RISSOINA

175

Distribution. Thus far Rissoina (Rissoina)
boucheti sp. nov. is only reported from Southern
New Caledonia.

Rissoina (Rissoina) longispira sp. nov.

Fig. 3 d-f

Type matériau. Ail from Biocal. stn DW 33,
29.VI1I.I985. Holotype (Fig. 3 d) : empty shell in
mnhn. Paratypes : 2 adult and one immature
empty shell in mnhn. Immature paratype coated
with gold for SEM-photography (Fig. 3 e-f).

Type locality. New Caledonia, 23°10' S,
167°10' E, 675-680 m.

Matériel examined. The type material is the
only one available.

Description. Shell : moderately large for
genus, very strongly elongate, conical.

Protoconch (Fig. 3 0 : dome-shaped, of 1 1/2
smooth whorls, of non-planktotrophic larval
type ; transition to teleoconch inconspicuous.

Teleoconch : of about 8 1/2, weakly convex to
almost straight-sided whorls ; sutures linear, very
weakly impressed.

Sculpture of very weak, irregular and rather
distantly spaced, weakly opisthocline, narrow,
axial ribs ; the latter becoming gradually less
regularly spaced and less prominent on penulti-
mate and last whorl ; axial ribs very weak to
almost absent on abapical half of last whorl ;
spiral sculpture of very weak, irregular and very
irregularly spaced spiral threads, on the abapical
half of spire whorls ; spiral threads slightly more
prominent on last whorl.

Aperture : D-shaped ; inner lip thin with a very
weak thickening near the transition to the ante-
rior channel ; the latter narrow, moderately
deep ; outer lip weakly thickened intemally, with
a weak, narrow varix externally ; outer lip weakly
opisthocline to almost orthocline in profile.

Colour : glossy white throughout.

Operculum : unknown.

Raduta and internai anatomy : unknown.
Shell dimensions : See table 2.

Table 2. — Rissoina <Rissoina) longispira sp. nov. Shell
dimensions. (L : shell length : i.s length of spire ; n : shell
diameter).

L (mm) Ls (mm) D (mm) No. whorls

Holotype (mnhn)
(Fig. 3 d)

9.7

6.9

3.0

8 1/2

Paratypes (mnhn)

9.4

4.1

3.2

8

8.8

6.3

2.8

7 3/4

Immature spccimcn
(Fig. 3 e-0

8.1

5.8

2.3

7 1/4

Variation. There is barely any variation in
shell size and sculpture in the small sériés of
specimens examined.

Etymology. — Longus (Latin) = long, spira
(Latin) = spire : referring to the strongly elon¬
gate spire.

Discussion. — Rissoina (Rissoina) longispira
sp. nov. is superficially similar to R. (R.)
boucheti sp. nov., but differs in having a less
angulate last whorl and in lacking the thick
swelling on the inner side of the outer lip, near
the transition to the anterior channel.

R. (R.) longispira is most similar to Rissoina
jaffa Cotton. 1952 from Cape Jaffa (South
Australia) and collected from a depth of about
500 m ; the latter species, however differs from R.
longispira in being much more acuminate, in
having a rather angulate last whorl instead of a
rather subglobose last whorl in R. longispira and
in having more prominent and somewhat more
opisthocline axial ribs. There are no other similar
species descri bed.

Distribution. — R. (R.) longispira sp. nov. is
thus far only known from the type locality.

Source : MNHN, Paris

176

WILLY J. SLEURS

Fig. 11. — Mantle cavity structure and anterior part of stomach of Rissoina (Rissoina) boucheli sp. nov., New Caledonia.
Biocal stn DW 44.

(Abbreviations : ac anterior chamber of stomach ; etc ctenidium ; k kidney ; oes oesophagus ; osphr
osphradium ; pe pericardium).

Genus ZEBINA H. & A. Adams, 1854

Zehina (Zebina) réclina sp. nov.

Fig. 4c-h

Type materiai.. — Ail from Biocal, stn DW 44,
30.VI1I.1985. Holotype (Fig. 4e-f) : empty shcll
in mhnh. Paratypes : 3 empty shells in mnhn.

Type locality. New Caledonia, 22“47' S,
167° 14' E, 440-450 m.

Material examined. The type materiai is the
only one available.

Description. Shell : moderately large for
genus, glossy, strongly elongate-conical.

Protoconch : of planctotrophic larval type,
of 2 1/4, slightly convex, weakly ad-abapically
compressed, smooth, whorls ; transition to teleo-

Source : MNHN, Paris

MOLLUSCA GASTROPODA : DEEP-WATER RISSOINA

177

conch abruptly, with a rather deep sinusigeral
notch.

Teleoconch : of about 7 to 7 3/4 whorls ;
adapical spire whorls weakly convex ; abapical
spire whorls becoming gradually less convex to
almost flat-sided ; last whorl moderately angu-
late ; suture linear, not impressed. Whorls smooth,
except for some very weak, densely spaced,
growth lines.

Aperture : pyriform ; inner lip thin posteriorly,
but becoming wider and thicker anteriorly near
the shell base ; anterior channel absent ; poste-
rior channel short, very narrow, triangular ;
outer lip thin except for a weak thickening near
the transition to the posterior channel and with
basal part strongly protracted ; outer lip exter-
nally with a moderately thick, rounded, narrow
varix.

Colour : glossy white throughout.

Operculum : unknown.

Radula and internai anatomy : unknown.

Shell dimensions : See table 3.

Table 3. /.china (Zebina) réclina sp. nov. Shell dimen¬
sions. (L : shell length ; i.s length of spire : d : shell
diameter).

L (mm)

Ls (mm)

D (mm) No. whorls

Holotype (mnhn) 5.9

3.8

2.3 7 3/4

Paratypes (mniin) 6.0

4.0

2.3 7 3/4

5.5

3.5

2.2 7 1/4

5.3

3.4

2.1 7

Variation. - The four specimens examined
(the type material) show only small différences in
shell dimensions, but do not dilTer in size and
shape.

Etymology. Reclinus (Latin) = leaning

back, referring to the strongly opisthocline outer
apertural lip.

Discussion. Zebina réclina sp. nov. is

superficially similar to Zebina acicula Laseron,
1956 from Christmas I. in shell shape, but difîers
in having a more angulate last whorl and in the
base of the outer lip being much more protrac¬
ted.

Z. réclina is easily distinguished from ail other

known congeners by the strongly angulate last
whorl.

Distribution. Zebina réclina sp. nov. is
known only from the type locality.

Zebina (Zebina) refusa sp. nov.

Fig. 4a-d

Type material. — Ail from Biocal. stn DW 38,
30.VIII. 1985. Holotype (Fig. 4 a-b) : empty shell
in mnhn. Paratypes : 5 adult. empty shells and
1 immature, empty shell in mnhn.

Type locality. — New Caledonia, 23°00' S,
167° 15' E, 360 m.

Material examined (Apart from type material).
New Caledonia. Biocal : stn DW 44. 22’AT S, 167" 14' E,
440-450 m, 30.V1II.1985 : I spec. (mnhn).

Description. — Shell : small, glossy. conical,
with bluntly rounded apex.

Protoconch : of non-planktotrophic larval
type, of about 2, relatively wide, smooth whorls ;
transition teleoconch abruptly.

Teleoconch : of 5 1/2 to 6 whorls; adapical
spire whorls slightly convex ; abapical spire
whorls becoming gradually less convex to almost
flat-sided ; last whorl subangulate near the peri-
phery.

Spire whorls and last whorl smooth, apart
from some weak growth lines.

Aperture : pyriform ; inner lip thin, becoming
moderately expanded anteriorly, partly covering
the shell base ; anterior channel absent : outer lip
thin, with about 3 weak parallel threads on the
outer margin of the inner side ; outer lip with a
moderately thick, rounded, narrow varix exter-
nally ; outer lip strongly opisthocline in profile.

Colour : spire whorls opaque white with some
very irregular large semitransparent dots just
above the suture ; last whorl with a rather wide
opaque band below the suture and with abapical
half semitransparent with irregular and irregu-
larly distributed subcircular opaque white dots.

Operculum, radula and internai anatomy :
unknown.

Shell dimensions : See table 4.

Source : MNHN, Paris

178

WILLY J. SLEURS

Table 4. Zebina (Zebina) retusa sp. nov, Shell dimen¬
sions. (i. : shell length : ls length of spire: d : shell
diameter).

L (mm) Ls (mm) D (mm) No. whorls

Holotype (mnhn)

3.4

2.3

1.4

5 3/4

(Fig. 4 a. b)
Paratypes (mnhn)

3.5

2.2

1.4

6

3.4

2.1

1.3

3.4

2.2

1.3

5 3/4

3.4

2.1

1.4

51/2

(Fig. 4c. d)

3.3

2.2

1.3

6

New Calcdonia

3.6

2.3

1.5

(Stn DW 44) (MNHN)

Variation. — Zebina retusa sp. nov. appears
to be very uniform in shell shape and shell size.

Etymology. Retusus (Latin) = blunt,
referring to the blunt apex of the shell.

Discussion. Zebina retusa sp. nov. difTers
from Z. réclina sp. nov. in having a bluntly
rounded apex instead of being strongly acumi-
nate, in the last whorl being less angulate and in
colour pattern.

Z. retusa difTers from ail its known congeners
in colour pattern and in the blunt, almost
rounded apex.

Distribution. Z. retusa sp. nov. is known
only from the type locality.

ACKNOWLEDGEM ENTS

I would particularly like to thank Dr P. Bou¬
chet, who provided me with the collection of
New Caledonian rissoinids.

I am very grateful to Dr G. Davis and to
Dr W. Ponder for their lessons on dissecting
techniques.

I would like to thank in particular Dr J. van
Goethem and Dr W. F. Ponder for providing
constructive comments on the original manus-
cript and for constant encouragement of my
research.

I wish to thank the following persons who
kindly sent me type material on loan : Mrs
K. Way (bmnh), Mrs Gowlett-Hoi.mes (South
Australian Muséum) and Mr P. Colman and
Mr I. Loch (ams).

The illustrations were rendered by Mr H. van
Paesschen (kbin), who also mounted the plates.
sem photography was done by Mr J. Cillis
(kbin).

REFERENCES

Cotton, B. C., 1952. - Australian Récent and
Tertiary Mollusca (Terebridae. Rissoinidae, Ris-
soidae, Fasciolariidae, Volutidae). Trans. R. Soc. S.
Aust.. 75 : 38-54, pl. 3-4.

Davis G.. Kitikoon V. & Temcharoen, P.. 1976. —
Monograph on " Lithoglyphopsis " aperta. the snail
host of Mékong River schistosomiasis. Malacologia,
15 (2) : 241-287, 22 figs.

Laseron, C. F., 1956. - The families Rissoinidae and
Rissoidae (Mollusca) from the Solanderian and
Dampierian zoogeographical provinces. Aust. J.
mar. Freshwat. Res., 7 (3) : 384-478.

Ponder. W. F., 1985. A Review of the Généra of
the Rissoidae (Mollusca : Mesogastropoda : Rissoa-
cea). Rec. Aust. Mus., Suppl. 4 : 1-221.

Powell, A. W. B., 1937. New species of marine
Mollusca from New Zealand. Diseovery Rep.. 15 ■
153-222. pl. 45-46.

Richer de Forges. B., 1990. Les campagnes
d'exploration de la faune bathyalc dans la zone
économique de la Nouvelle-Calédonie. /// : Résul¬
tats des campagnes Musorstom. Volume 6 (1).
Mém. Mus. nain. Hist. mit. Paris , (A) 145 : 9-54.

Source : MNHN, Paris

ILTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSULT

6

Mollusca Gastropoda : Cypraeopsis superstes sp. nov.,
Pediculariinae relique du bathyal
de Nouvelle-Calédonie et de la Réunion

Luc DOLIN

2, place du Caquel
93200 Saint-Denis
France

RÉSUMÉ

Le genre Cypraeopsis n'étail connu que par deux espèces
du Miocène d'Europe et d'Asie du Sud-Est. Une espèce non
dénommée est signalée ici de l'Oligocène supérieur de France
et C. superstes sp. nov. est décrite de la faune bathyale actuelle
de la Nouvelle-Calédonie et de la Réunion. C. superstes

diffère des espèces fossiles par la sculpture finement côtelée
du dernier tour et le bord externe ondoyant, annonçant les
Pedicularia , par sa protoconque saillante, non ennoyée un
caractère peut-être progènétique . C. superstes apparaît,
paradoxalement, comme une relique évoluée.

ABSTRACT

Cypraeopsis superstes sp. nov., a ne» bathyal relict spccics
of Pediculariinae (Gastropoda Ovulidae) front New Caledonia
and Réunion.

The genus Cypraeopsis was so far known from two species
in the Miocène of Europe and South-East Asia. An unnamed
species is here recorded from the upper Oligocène of France

and C. superstes sp. nov. is described from the Recent
bathyal fauna of New Caledonia and Réunion. C. superstes
differs from the fossil species by the body whorl being spirally
sculpturcd. by the outer lip undulating as in Pedicularia. and
by the protruding. uncovered protoconch. a character tenta-
tively interpreted as progenetic. C. superstes thus appears
paradoxically as an evolved relict.

Dolin, L.. 1991. Mollusca Gastropoda : Cypraeopsis superstes sp. nov.. Pediculariinae relique du bathyal de Nouvelle-Calédonie et de
la Réunion. In : A, Crosnikr & P. BOUCHET (eds). Résultats des Campagnes Musorstom, Volume 7. Ment. Mus. nain. Hist. nat.. (A). 150 ;
179-186. Paris ISBN : 2-85653-180-6.

Publié le 20 mars 1991.

Source : MNHN, Paris

180

LUC DOLIN

INTRODUCTION

Cette étude repose sur le matériel récolté dans
le sud de la Nouvelle-Calédonie (voir Richer de
Forges, 1990) et à la Réunion, lors de campagnes
des navires océanographiques “ Jean Charcot ”,
“ Vauban ”, “ Coriolis " et “ Marion Dufresne ”,
La faune bathyale de Nouvelle-Calédonie pré¬

sente une richesse remarquable en Stylastérides
(plusieurs dizaines d'espèces). C. superstes y est
sympatrique avec Pedicularia (s. s.) cf. elegantis-
sima Deshayes, 1863, qui a été récolté sur son
hôte (Stylasteridae).

ÉTUDE SYSTÉMATIQUE
Superfamille CYPRAEOIDEA Rafinesque, 1815

Famille OVULIDAE Fleming, 1822
Sous-Famille PEDICULARIINAE Gray, 1853

Jusqu'à ce que Gosliner et Liltved (1985 :
105, 118-119, fig. 34) aient établi sur la base de
critère anatomiques l’appartenance des Pedicula-
riinae aux Ovulidae, cette entité phylétique a
maintes fois changé et de rang, et de contenu.
Schilder (in Wenz, 1941 : 1000-1003, 1007-
1008) intégrait notamment au sein des Pedicula-
riinae les genres monospécifiques éocènes Semi-
cypraea Schilder, 1936 (= Cypraeidae, Ber-
nayinae ; voir Dolin & Dolin, 1983 : 18-19,
fig. 5 a-b), Cypraeogemmula Vredenburg, 1920
(= Ovulidae, Sulcocypraeinae) et Eovolva Schil¬
der. 1932; mais alors que les Cypraedia Swain-
son, 1840 y étaient bien associées aux Pedicularia
Swainson, 1840, les Jenneria Jousseaume, 1884 et
Cypraeopsis Schilder, 1936 étaient relégués au

sein des Sulcocypraeinae (= Eocypraeinae). Or
Cypraedia (voir Martin, 1914 : fig. 121b),
Cypraeopsis (Fig. 2 a-b) et Pedicularia (Schilder
1931 ; fig. I ; Richter & Thorson, 1975 : pl. 9,
fig. 58 a-b) possèdent une protoconque en tous
points semblables, sinusigériforme, multispirée à
sculpture obliquement treillissée (décussée), et
non verticalement quadrillée comme chez les
Cypraeidae (voir Thiriot-Quievreux, 1967 : pl.
1-3 ; Richter & Thorson, 1975 ; pl. 8, lig. 52-
57 ; pl. 9. fig. 59-60 ; Dolin & Dolin, 1983 : fig.
3-4, 25, 26-b). De par ses spécificités originales,
C. superstes permet par ailleurs de situer les
Cypraeopsis à mi-chemin des Jenneria (préda¬
teurs de Scléractinaires) et des Pedicularia (pré¬
dateurs d’Hydrozoaires).

Genre CYPRAEOPSIS Schilder, 1936

Cypraeopsis Schilder, 1936 : 87. Espèce-type, par
désignation originale : Cypraeopsis vandervlerki Schil¬
der. 1936.

« ... allied to Transovula in size, in general
shapc, and in the dentition of both lips, but
differs by being less elongate, by the longitudinal

dorsal sulcus, by a longitudinal callous carina, by
the réduction for any thickening of the inner lip
on its posterior extremity. and by the distinctly
denticulate fossula ».

C. superstes diffère essentiellement de ses
ancêtres par la structure finement côtelée de son

Source : MNHN, Paris

MOLLUSCA GASTROPODA : CYPRAEOPS/S DU BATHYAL

181

aire dorsale, par son canal anal peu marqué et
par sa lèvre externe ondoyante. Je n'ai toutefois
pas cru devoir sanctionner taxonomiquement
cette évolution morphologique. Elle pourrait en
effet être de nature adaptative : bien qu’aucun
échantillon de C. superstes n'ait été récolté avec
son hôte, le contour de la lèvre externe pourrait
être induit par la forme d’une branche de
Stylasteride, par exemple, comme c'est le cas
chez les Pedicularia (Arnaud & Zibrowius,
1979 : 123-124; Schmieder, 1982 ; 272, fig. 1).

Cypraeopsis superstes sp. nov.

Fig. 1 a-c, 2 a-b, 3 a-b, 4.

Matériel type. — Holotype mnhn (Smib 3,
st. DW 12 : Fig. 1 a-c, 4) et paratypes mnhn
(Bioc al, si. DW 51 : Fig. 2 a-b. — Musorstom 4,
st. DW 197 : Fig 3 a-b) ; paratypes nmnz, ams,
nsmt, et nm (Musorstom 4, st. DW 197).

Localité type . Smib 3, st. DW 12,
23°38' S-167°42' E, 470 m.

Matériel examiné. Nouvelle-Calédonie. Bio¬
cal. coll. Bouchet, Metivier et Richer de Forges :
St. DW 08. 20‘34'S, 166°54'E, 435 m : 13 spms
(population naine, adulte ne dépassant pas 6 mm). —
St. DW 33, 23° 10' S. 167°10' E. 675-680 m : 3 spms.

St. DW 48. 23°00' S, 167°29' E, 775 m : 20 spms. —
St. DW 51, 23°05' S, 167°45' E. 680-700 m : 109 spms
+ récoltés vivant (nombreux stades juvéniles). —
St. DW 53. 23°09' S. 167°43' E. 975-1 005 m : 1 spm.
St. DW 66, 24°55' S. 168°22' E. 515-606 m : 1 spm.
St. DW 70, 23°25' S. 167°53' E. 965 m : 3 spms.

St. DW 77. 22° 15'S, 167° 15' E. 440 m : 1 spm.

St. DW 83. 20°35'S, 166°54' E. 460 m : 2 spms.

Musorstom 4, coll. Bouchet et Richer de Forges :
St. DW 156. 18°54'S. 163° 19' E, 530 m : 2 spms +
vivant. St. DW 159, 18°46' S, 163° 16' E. 600 m ; 3
spms. St. DW 162, 18°35'S. 163°10'E, 535 m 1
spm, — Si. CP 194, 18°53' S, 163°22' E. 550 m : 1 spm
+ 2 spms récoltés vivants. St. DW 197. 18°51'S.
163°21’ E, 560 m : 13 spms + 10 spms récoltés vivants.

St. DW 220, 22°58' S. 167°38' E. 505-550 m : 2 spms.
— St. CP 216. 22°59' S. 167°22' E. 490-515 m : 2 spms
récoltés vivants. - St. DW 221. 22°59'S. 167°37' E,
535-560 m : 2 spms. — St. DW 223, 22°57' S.
167°30' E. 545-560 m : I spm. St. DW 225. 22°52' S.
167°23' E. 590-600 m : 1 spm.

Chalcal 2, coll. Bouchet. Metivier et Richer de
Forges : St. DW 72, 24°55' S, 168°22'E. 527 m :
1 spm. St. DW 76, 23°41'S, 167°45' E, 470 m :
8 spm.

Smib 3. coll. Richer de Forges : St. DW 12,

23°38'S, 167°42'E. 470 m : 1 spm. — St. DW 24.
22°59'S, 167°21'E, 535 m : 4 spms.

Réunion : Campagne MD 32 : St. DC 10, 21 ° 13' S,
55°52' E. 930-980 m : 1 spm + 6 spms juvéniles.

Description. — Coquille piriforme, à test
mince et à lèvre ondoyante, contournant la zone
apicale où elle se confond avec l'arête adapicale
du bord interne. L'aire dorsale et la denticulation
sont blanches. Une auréole beige pâle, passant
graduellement au marron-rouge, souligne les
marges latérales et les extrémités. L'unique spéci¬
men adulte de la Réunion montre une proto-
conque brune, mais la téléoconque est d'un blanc
immaculé.

Protoconque brun-rouge, en partie ennoyée
dans le dernier tour de téléconque, faisant légè¬
rement saillie au creux d’une dépression pseu-
dombilicale. Sur les exemplaires juvéniles, on
distingue une protoconque I lisse, avec seulement
quelques granules suprasuturaux. Lui succède
une protoconque II composée de 3 tours envi¬
ron, à sculpture formée de croisillons obliques
(Fig. 2 a-b). La transition protoconque/téléo¬
conque (très distincte lorsqu'elle n'est pas recou¬
verte par le tour suivant) montre la lèvre à 3
échancrures caractéristiques de la superfamille.

L'aire dorsale est parcourue d'une multitude
de côtes spirales émoussées, équidistantes (6 à 7
au mm en moyenne), séparées par des sillons
profondément incisés. De fines stries d'accroisse¬
ment les recoupent plus ou moins régulièrement.
Le canal siphonal est remarquablement allongé,
assez profond bien délimité ; bien détaché, il
forme un cou net. Le canal anal est beaucoup
plus mal formé. Il n'est que faiblement limité
abaxialement par les 3 dernières fortes dents de
l’arête adapicale, et ne se creuse d’aucune gout¬
tière. Le bord interne est fortement convexe,
formant une crête longitudinale calleuse, forte¬
ment versante adaperturalement. Trente à qua¬
rante fortes rides parcourent l'aire basale divisée,
ainsi, en deux versants ; se dédoublant parfois,
elles sont atfaiblies au niveau du versant adaper-
tural, affaissé, concave en sa partie médiane. Le
pli antérieur horizontal, remarquablement pro¬
longé, et sa lame interne raccourcie à l'extrême,
sont réduits à leur plus simple expression. La
fossula est résiduelle, légèrement concave, et
l'aire columellaire est étroite ; elles sont irréguliè¬
rement denticulées. Lèvre externe ondoyante à
marge lamelleuse, bordée, formant un méplat
que parcourent une quinzaine de fortes dents

Source : MNHN, Paris

182

LUC DOLIN

Source : MNHN, Paris

MOLLUSCA GASTROPODA : CYPRAEOPS1S DU BATHYAL

183

faisant saillie sur l’ouverture ; de fines rides
s'intercalent entre elles, limitées à l’extérieur du
méplat et sans solution de continuité avec les
côtes et sillons de faire dorsale. Le développe¬
ment adapicale en pavillon de la lèvre externe
accuse la largeur de l’ouverture.

La radula est taenioglosse (Fig. 3 a-b) : dent
centrale avec une large plaque basale, un fort
cuspide médian et 3-4 cuspides plus petits de
chaque côté ; dent latérale avec un très long et
fort cuspide et, respectivement, 1 et 2-3 cuspides
vers l’intérieur et l’extérieur de la dent ; margina¬

les longues, la dent interne étant emboîtée par sa
base dans la dent externe.

Distribution. Bathyal du Sud de la Nou¬
velle-Calédonie et de la Réunion, les exemplaires
vivants ayant été pris en Nouvelle-Calédonie
entre 490 m et 700 m. Une plus vaste distribution
Indo-Ouest Pacifique est envisageable.

Etymologie . — Pour qualifier l'unique repré¬
sentant actuel d'un genre Oligo-Miocène que
l'on croyait éteint, superstes — le survivant —
s’imposait.

Discussion

Affinités : Le premier Cypraeopsis est connu avec
certitude du Chattien (Oligocène supérieur, 26 mil¬
lions d'années environ). L'unique spécimen (Fig. 4)
a été récolté à « Estoti », Saint-Paul-lès-Dax
(Landes, France) dans le faciès de base dont la
thanatocoenose « indicates sédimentation on the
lower beach environment » (Lozouet in Dolin
et al., 1985 : 10). Cette espèce, vraisemblable¬
ment nouvelle, annonce très précisément l’espèce
suivante.

C. subursellus (d’Orbigny, 1852) du Burdiga-
lien (Miocène inférieur) du bassin d’Aquitaine
(France ; fig. 5) et des collines de Turin (Italie ;
voir Ferrero Mortara et al., 1984 : pl. 26,
fig. 6 a-b) en est toutefois spécifiquement distinct.
En moyenne que son ancêtre, C. subursellus
présente une denticulation du bord interne plus
dense, tandis que celle du bord externe est
dédoublé comme chez C. superstes ; par ailleurs,
la fossula paraît bien avoir amorcé une réduction
sensible, la denticulation s’accentuant au péris-
tome de la fossula. Le C. sp. de l'Oligocène
supérieur et C. subursellus se rencontrent dans
des milieux sensiblement similaires, de récifs
coralliens plus ou moins démantelés ; ils y sont
systématiquement associés à des espèces du
genres voisin Jenneria.

C. vandervlerki des « Taballar Kalk » (Mio¬
cène supérieur?) de Bornéo (Indonésie), est
fondé sur un unique exemplaire figuré dans
Wenz (1941 : fig. 2894). Cylindracée, elle se
distingue nettement des deux espèces européennes
par sa denticulation réduite, y compris sur la
fossula. Pour le reste, elle montre les caractères
génériques — notamment — du pli antérieur

court, des deux bords carénés, etc., jusqu'à et y
compris la sculpture du juvénile qui persiste chez
les trois espèces fossiles au niveau de faire
columellaire (et que l'on pourrait prendre pour
une denticulation interne).

En dépit d'un changement d'environnement
radical (phénomène pour lequel je n'ai pas d'expli¬
cation), et d'un écart stratigraphique avoisinant
vraisemblablement une quinzaine de millions
d’années, C. superstes présente nombre de carac¬
tères ancestraux, inchangés ou si peu. Elle offre
même avec C. subursellus beaucoup d'affinités :
le pli antérieur (davantage encore prolongé) : la
lame interne (quoique plus réduite) : la denticula¬
tion forte du bord interne, à cheval sur la crête
longitudinale affectant faire ventrale, légèrement
affaissée : le bord externe formant un méplat,
bordé (devenant lamelleux). fortement denticulé,
des rides s'intercalant entre les rides principales
(limitées au péristome extérieur du labre).
Caractères taxonomiques : C. superstes ne diffère
donc vraiment de ses ancêtres que par des
détails, importants et rappelant certains genres,
mais que je pense être d'ordre spécifique.

1. Alors que la protoconque est chez les
espèces fossiles ennoyée par une épaisse callosité
couronnant le canal anal, chez C. superstes son
originalité est de se situer — même au stade
adulte — au fond d'une dépression pseudombili-
cale, et de n’être jamais convolutée par le dernier
tour — comme c’est le cas pour la quasi totalité
des Ovulidae (voir Ranson, 1967 : 97-98, fig.
144-145, 148-145, 148-149). Ceci pourrait indi¬
quer une progénèse.

2. Les Cypraeopsis fossiles, comme les Jenneria.

Source : MNHN, Paris

LUC DOLIN

184

Figs 4-7. — 4. Cypraeopsis superstes, hololype mnhn, smib 3 st. DW 12 (Nouvelle-Calédonie), 11,8 mm. 5, Cypraeopsis sp
Estoii, Saint-Paul-lés-Dax (Landes, France), collection A. Cluzaud. Chaltien (Oligocène supérieur), 13,6 mm. 6
Cypraeopsis subursellus (d'Orbigny), 1852). Moulin de Carreau. Corbieux (Landes, France), collection R. Favia'
Burdigalien (Miocène inférieur sommital), 20.4 mm. 7. Cypraeopsis vanaervlerki (d'après Wenz, 1941). Bornéo
(Indonésie). Taballar kalk (Miocène supérieur?), 9,4 mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : CYPRAEOPSIS DU BATHYAL

185

possèdent un sulcus dorsal, accentuation de la
zone de jointoiement des deux lobes du manteau
par l'hypertrophie des cals du canal siphonal, du
canal anal et des marges. Chez C. superstes,
l’aire dorsale quoiqu'épaisse — est finement
côtelée, ces costules spirales étant recoupées par
les stries d'accroissement. Cette structure ne se
retrouve telle quelle que chez les Pedicularia, et
tout particulièrement chez Pedicularia (Pedicula-
riella) californica (Newcomb, 1864).

3. Bien que les caractères du bord interne et
de sa denticulation soient identiques chez C.
subursellus et C. superstes, chez cette dernière la
frange labrale ondule, se développant postérieu¬
rement en pavillon, sans former ni délimiter de
canal exhalant. Ce critère préfigure également de
manière évidente les Pedicularia, chez qui cette
aponiorphie correspond à une adaptation au
commensalisme sur des Hydrocoralliaires Stylas-
terides (Arnaud & Zibrowius, 1979 : 123-124;
Schmieder, 1982 : 272, fig. 1).

Somme toute, si les Cypraeopsis fossiles se
rattachent morphologiquement aux Jenneria, avec

lesquels ils ont — au moins au début partagé
le même biotope, C. superstes fait le lien et
annonce les Pedicularia ; aucune coupure intra-
générique ne peut toutefois être sérieusement
établie au sein des Cypraeopsis. Un rattachement
aux Pedicularia ne serait pas plus fondé.

En effet, Pedicularia (s.s) cf. elegantissima
Deshayes, 1863 avec laquelle C. superstes est
sympatrique, montre (Biocal. st. DW 57 ; à
publier) une radula identique à celle figurée par
Fischer (1887 : 665, fig. 423) reprise par Thiele
(1929 : 269, fig. 284), figures en partie erronées
car montrant une marginale en forme d'écaille,
qui n’existe pas (Bouchet, comm. pers.) ; or,
celle de C. superstes en diffère notablement par
sa dent centrale à cuspides moins nombreux mais
longs et acérés, détachés les uns des autres. Sa
radula lui confère donc une aussi grande origina¬
lité que les caractères morphologiques de son
test, situant les Cypraeopsis à leur place, au sein
des Pediculariinae, à mi-chemin des Jenneria et
des Pedicularia.

CONCLUSION

Unique représentant vivant du genre Cypraeo- caractéristiques qui conduisent à la qualifier
psis, avec qui plus est une distribution paradoxalement de « relique évoluée ».
bathyale, C. superstes présente un ensemble de

REMERCIEMENTS

Le matériel actuel étudié a été récolté lors de
campagnes conjointes orstom/mnhn ; je vou¬
drais associer aux collecteurs cités, Virginie
Héros et Annie Tillier (mnhn) pour leur travail
patient et leur aide concrète.

Les échantillons fossiles proviennent de fouilles
individuelles ; Raymond Favia et Alain Cluzaud
les ont amicalement mis à ma disposition.

Anders Warèn du Naturhistorika Riksmuseet,

Stockholm, a aimablement distrait de son temps
pour extraire les radula de Cypraeopsis superstes et
Pedicularia cf. elegantissima. Les photographies
sont de Pierre Lozouet et de Philippe Bouchet
(meb), qui a, en outre, corrigé les manuscrits
successifs.

Que tous trouvent ici l’expression sincère de ma
reconnaissance.

Source : MNHN, Paris

186

LUC DOLIN

RÉFÉRENCES BIBLIOGRAPHIQUES

Arnaud, P. M. & Zibrowius, H., 1979. L’associa¬
tion Pedicularia sicula Errina aspera en Mediter¬
ranée (Gastropoda Prosobranchia et Hydrocorallia
Stylasterina). Rapp. Comm. int. Mer Médit., 25/26
(4) : 125-126.

Dolin. C. & Dolin, L., 1983. Révision des Tri-
viacea et Cypraeacea (Mollusca, Prosobranchiata)
éocènes récoltés dans les localités de Gan (Tuilerie
et Acot) et Bosdarros (Pyrénées Atlantiques, France).
Meded. Werkgr. Tert. Kwart. Geol., (1) 20 : 5-48.

Dolin. C„ Dolin. L. & Lozouet, P.. 1985. —
Paleoecology of some classic Tertiary localities in
the Aquitaine and Paris basins of France. Mississipi
Geology, 5 (4) : 4-13.

Ferrero Mortara. E.. Montefameglio, L.. No-
VELLI, M.. Opesso. G.. Pavia. G.. Tampieri, R.,
1984. Catalogo dei tipi c degli esemplari figurati
délia collezione Bellardi e Sacco. Cataloghi Mus.
reg. Sci. nat. Torino. 7 (2) : 1-484.

Fischer, P.. 1887. — Manuel de Conchyliologie. F.
Savy. Paris : 1-1369, pis 1-23.

Gosliner. T. M. & Liltved. W. R.. 1985. — Aspects
of the morphology of the endemic South African
Cypraeidae with a discussion of the évolution of the
Cypraeacea and Lamellariacea. Ann. S. Afr. Mus..
96 (4) : 67-122.

Martin. K.. 1914. Die Fauna des Obereocâns von
Nanggulan, auf Java. Samml. Geol. Reichmus. Lei¬
den. (n.s.) 2 : 110-223, pis 1-8.

Ranson. G., 1967. — Les protoconques ou coquilles
larvaires des Cyprées. Mëtn. Mus. nain. hist. nat.
(A) 47 (2) : 93-126, pl. 1-39.

Richer de Forges, B., 1990. Les campagnes
d'exploration de la faune bathyale dans la zone
économique de la Nouvelle-Calédonie. In A. Cros-
nier (éd.). Résultats des campagnes Musorstom,
Volume 6. Mém. Mus. nain. Hist. nat., (A). 145 : 9-
54.

Richter. G. & Thorson. G.. 1975. — Pelagischc
Prosobranchier-larven des Golfes von Neapel. Oplie-
lia. 13 : 109-185.

Schilder, F. A.. 1931. Révision of the family
Pediculariinae. J. Conch.. 19 : 165-169, pl. 6.

Schilder, F. A.. 1936. — Anatomical characters of
Cypraeacea which confirm the conchological clas¬
sification. Proc. Mal. Soc. Lond.. 22 (2) : 75-112.
pl. 11-12.

Schmieder R. W., 1982. — Shape irregularity in
Pedicularia californica. Veliger, 24 (3) : 272. fig. 1.

Thiele, J.. 1929. — Handbuch der Systematischen
Weichtierkunde, 1 (1). G. Fischer, Jena : 1-376.

Thiriot-Quievreux. C.. 1967. — Observations sur le
développement larvaire et postlarvaire de Simnia
spelta Linné (Gastéropode Cypraeidae). Vie & Milieu.
18 ( 1 -A) : 143-151.

Wenz. W.. 1941. — Handbuch der Palâozoologie, 6,
(5). O. H. Schindewolf, Berlin : 949-1014 (Cypraea¬
cea).

Source : MNHN, Paris

ILTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSUL

7

Mollusca Gastropoda :

from New

On a collection of Nassariidae
Caledonian waters

Walter O. CERNOHORSKY

6 Rapallo Place
Farm Cove. Pakuranga
Auckland 6
New Zealand

ABSTRACT

The présent report deals with a collection of 33 species of
Nassariidae from New Caledonian waters. Approximately
30 % of the species recorded are new geographical range
extensions. Nassarius bifarius (Baird in Brenchley. 1873),
previously considered a synonym of N. novaezelandiae
(Reeve. 1854). and N. stigmarius (A. Adams. 1852), pre¬

viously considered a synonym of N. splendidulus (Dunkcr.
1846). arc no» acknowlcdged to be valid. separate species.
Nassarius nlomea Kay. 1979 is synonymized with N. crebri-
ctislalus (Schcpman. 1911). Nassarius iZeuxls) arcus sp. nov.
is described and recorded from depths of 95-200 m.

RÉSUMÉ

Mollusca Gastropoda : Sur une collection de Nassariidae des
eaux néo-calcdonicnnes.

Trente-trois espèces de Nassariidae sont signalées de la
région néo-calédonienne, dont un tiers pour la première fois.
Ce matériel permet de reconnaître la validité de Nassarius
bifarius (Baird in Brenchley, 1873). jusqu'ici traité comme

synonyme de N. novaezelandiae (Reeve. 1854). et N. stigma¬
rius (A. Adams. 1852), jusqu'ici traite comme synonyme de
N. splendidutus (Dunkcr, IK46). Nassarius olomea Ka>. 1979
est placé en synonymie de N. erebricostalus (Schcpman.
1911). Nassarius (Zeuxist arcus sp. nov. est décrit d'après
des spécimens péchés entre 95 et 200 m de profondeur

Cernohorsky, W. O.. 1991. - Mollusca Gastropoda : On a collection of Nassariidae from Ne» Caledonian waters In A Ckusmi r &
P. Bouchet (eds). Résultats des Campagnes Musorstom, Volume 7. Mcm Mus. nain. Mise nat.. (A). 150 : 187-204 Pans ISBN 2-85653-180-6
Publié le 20 mars 1991.

Source : MNHN, Paris

188

WALTER O. CERNOHORSKY

INTRODUCTION

Through the courtesy of Dr P. Bouchet,
Department of Malacology, Muséum national
d’Histoire naturelle, Paris, I hâve received for
détermination material of Nassariidae brought
together by various expéditions and dredging
programs in New Caledonian and adjacent waters.
There are two main sources of material. One is a
sériés of deep-sea expéditions in the Chesterfield
islands and New Caledonia. For a narrative of
these cruises, including complété station lists, see
Richer de Forges (1990). The other is the
“ programme Lagon ” conducted for Orstom
since 1984 by Dr B. Richer de Forges aboard
R. V. “ Vauban " and R. V. "Alis " in the New

Caledonian coral reef lagoon. This program
comprises over 1000 dredging and trawling sta¬
tions operated on a 2 nautical miles grid, mostly
in depths between 10 and 60 m, with a few hauls
in deeper water. In the présent report, stations
hâve been grouped according to the major
geographical divisions of the New Caledonian
lagoon. Full station data are to be found in
Richer de Forges & Bargibant (1985) and
Richer de Forges et al. (1987).

Species are arranged alphabetically within
généra and subgenera, and the more significant
species illustrated. The abbreviation spm refers
to live-taken specimens as well as empty shells.

SYSTEMATIC ACCOUNT
Family NASSARIIDAE

Genus NASSARUJS Duméril, 1806

Nassarius (Alectrion) g laits glans (Linnaeus, 1758)

Buccmum glans Linnaeus, 1758 : 737.

Nassarius (Alectrion) glans glans - Cernohorsky,
1984 : 61, pl. 2, figs 5-7 (synonymy).

Material examined. New Caledonia. Lagon :
Surprise Atoll : stn 465, 45 m : 1 spm.
Southwestern Lagoon : stn 3, 15 m : 3 spm. Stn 4,
9 m : 8 spm. — Stn 5, 10 m : 2 spm. Stn 7, 14 m:
2 spm. — Stn 9. 10 m : 1 spm. — Stn 21, 10 m : 2 spm.

Stn 22. 11 m : 1 spm. Stn 23, 10-18 m : 1 spm.
Stn 25, 28 m : 1 spm. — Stn 34, 10 m : 1 spm.

Stn 38, 20 m : 1 spm. Stn 41, 28-46 m : 1 spm. —
Stn 49, 10 m : 1 spm. Stn 50, 12 m : 2 spm.

Stn 57, 10 m : 1 spm. Stn 60. 11m: 2 spm. —

Stn 66, 15 m : 3 spm. — Stn 67, 21 m : 2 spm.

Stn 79, 16 m : 1 spm. Stn 80, 33 m : 10 spm. —

Stn 83, 22 m : 7 spm. Stn 84. 17 m : 8 spm. —

Stn 99, 17 m : 4 spm. — Stn 100, 15 m : 4 spm. —

Stn 121, 12 m: 1 spm. — Stn 129, 44-55m : 1 spm. —
Stn 156, 21 m : 1 spm. — Stn 161, 20 m : 3 spm.

Stn 166, 10 m : 1 spm. Stn 169, 22 m : I spm. —

Stn 170, 22m : 1 spm. Stn 185, 15 m : 5 spm.

Stn 192, 18 m : 2 spm. Stn 201, 17 m : 4 spm. —

Stn 206, 8 m : 2 spm. - Stn 210, 14 m : 1 spm.

Stn 212, 10m : 1 spm. Stn 229, 41 m : I spm. —

Stn 233, 30m : 1 spm. — Stn 247, 43 m : 1 spm.

Stn 249. 11 m : 2 spm. Stn 250. 10 m : 1 spm. -

Stn 252, 22 m : 2 spm. - Stn 253, 16 m : 2 spm. —

Stn 259, 18 m : 3 spm. Stn 269, 20 m : I spm. —

Stn 272, 12m : 1 spm. — Stn 275, 19 m : I spm. -

Stn 282, 12 m : 1 spm. Stn 284, 6 m : 2 spm. —

Stn 293, 20 m : 2 spm. Stn 295, 41m: 1 spm. —

Stn 298, 37 m : I spm. Stn 305, 26 m : 1 spm. —

Stn 306, 38 m : I spm. — Stn 312, 26 m : 1 spm. —

Stn 339, 26 m : I spm. Stn 340, 27 m : 1 spm._

Stn 349, 55 m : 1 spm. - Stn 547, 29 m : 1 spm. _

Stn 548, 32 m : 1 spm. — Stn 550, 24 m : 3 spm. -

Stn 555, 32 m : 2 spm. Stn 591, 14 m : 5 spm. —

Eastern Lagoon : stn 623, 32-40 m : I spm. —

Stn 627, 45-47 m : 1 spm. — Stn 635, 51 m : 1 spm. —

Stn 671, 36-39 m : 1 spm. Stn 698, 40-43 m :
1 spm. — Stn 712, 47-49 m : I spm. Stn 716
30 m : 1 spm. — Stn 731, 37-42 m : 1 spm.

Distribution. From the Persian Gulf to

Japan and the Marquesas Islands.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NASSAR1IDAE

189

Nassarius ( Pliearcularia) fraudulentus
(Marrat, 1877)

Nassa frauda lent a Marrai, 1877 : 8, pl. 1, fig. 24.
Nassarius (Pliearcularia) fraudulentus - Cernohorsky,
1984 : 72, pl. 5, figs 11, 12 ; pl. 6, fig. 1 (synonymy).

Materjal examined. Chestcrfield Islands Chal-
cal I : stn DC 21, 19°18'S. 158“43'E, 73 m, 17 July
1984 : 2 spm.

New Caledonia. Lagon :

Southwestern Lagoon : Sln 14, 22 m : 1 spm. —
Stn 19, 29 m : 1 spm. Stn 31, 29 m : 1 spm. —
Stn 69, 13m : 1 spm. — Stn 78, 35m : 11 spm. —

Stn 86, 29 m : 3 spm. Stn 122, 28 m : 1 spm. —

Stn 167, 11 m : 2 spm. — Stn 174, 45 m : I spm. —

Stn 178, 20 m : 6 spm. Stn 188, 8m : 1 spm. —

Stn 192, 18 m : I spm. — Stn 193, 20 m : 2 spm. —

Stn 195, 12m : 1 spm. Stn 199. 50 m : 1 spm. —

Stn 203, 13 m : 8 spm. — Stn 205. 13 m : 1 spm.

Stn 206, 8 m : 1 spm. Stn 208, 9 m : 1 spm. —

Stn 209, 14 m : 5 spm. Stn 210, 14 m : 2 spm. —

Stn 211, 12 m : 8 spm. Stn 215, 14 m : 24 spm.

Stn 216, 14 m : 17 spm. Stn 219, 32 m : 6 spm.

Stn 220, 12 m: Il spm. — Stn 222. 24 m : 2 spm.

Stn 261, 19 m : 3 spm. Stn 274, 12 m : 3 spm. —

Stn 277, 12 m : 3 spm. — Stn 278, 17 m : 2 spm. —

Stn 287, 29 m : 1 spm.

Eastern Lagoon : stn 691, 33-34 m : 1 spm.

Distribution. - From Mauritius to India
and the Society Islands.

Nassarius (Pliearcularia) globosus
(Quoy & Gaimard, 1833)

Buccinum globosum Quoy & Gaimard, 1833 : 448,
pl. 32, figs 25-27.

Nassarius (Pliearcularia) globosus - Cernohorsky,
1984 : 74, pl. 6, figs 7-10 (synonymy).

Material examined. New Caledonia. Lagon :
Southwestern Lagoon : stn 221, 55-65 m : 5 spm.

Distribution. From the Red Sea to Japan
and the Samoa Islands.

Nassarius (Pliearcularia) granifer (Kiener, 1834)

Buccinum graniferum Kiener, 1834 : 100, pl. 27,
fig. 111.

Nassarius (Pliearcularia) granifer - Cernohorsky :
76, pl. 7, figs 1-2 (synonymy).

Mait-rial examined. New Caledonia. Lagon :
H non Atoll : stn 443, 40m : 1 spm.

Surprise Atoll : stn 449, 21 m : 3 spm.

Northern Lagoon : stn 481, 33 m : 2 spm.
Southwestern Lagoon : stn 8, 15m 1 spm. —

Stn 49, 10m : 2 spm. Stn 79, 16m : 1 spm.

Stn 98, 15 m : 2 spm. — Stn 159. 17 m : 4 spm.

Stn 162, 10 m : 2 spm. — Stn 214, 12m : 1 spm.

Stn 218, 15m : 1 spm. — Stn 283, 13 m : 3 spm.

Stn 308, 18 m : 4 spm. — Stn 549, 26 m : 1 spm. —

Stn 551, 9 m : 2 spm.

Distribution. — From East Africa through-
out the tropical Indo-Pacific to the Tuamotu
Archipelago.

Nassarius (Pliearcularia) troendleorum
Cernohorsky, 1980
Figs 1-2

Nassarius troendleorum Cernohorsky, 1980 : 118,
figs 15-17 (shell), fig. 25 (protoconch).

Nassarius (Pliearcularia) troendleorum - Cernohorsky,
1984 : 77, figs 107, 108.

Material examined. New Caledonia. Lagon :
Northern Lagoon : stn 489, 43 m : 1 spm.

Distribution. — From New Caledonia to the
Kingsmill Group, Gilbert Islands and the Tua¬
motu Archipelago.

Nassarius (Niotha) albescens albescens
(Dunker, 1846)

Buccinum albescens Dunker, 1846 : 170.

Nassarius (Niotha) albescens albescens - Cerno¬
horsky, 1984 : 81. pl. 7, figs 11, 12: pl. 8, fig. I
(synonymy).

Material examined. — New Caledonia. Lagon :
Surprise Atoll : stn 449. 21 m : 3 spm.
Southwestern Lagoon : stn 21, 10 m : 2 spm. — Stn 49.

10 m : 2 spm. — Stn 98. 15 m : 2 spm. — Stn 162.

10 m : 2 spm. — Stn 214, 12 m : 1 spm. Stn 296,

26 m : 1 spm. Stn 341. 19 m : 2 spm. Stn 554,

27 m : 4 spm.

Distribution. From the Nicobar Islands to
Japan and the Cook Islands.

Nassarius (Niotha) bifarius
(Baird in Brenchley, 1873)

Figs 3-6

Nassa bifaria Baird in Brenchley, 1873 : 436. pl. 38,
figs 1, 2 (Type locality : New Caledonia).
Nassarius (Niotha) bifarius - Cernohorsky, 1972 :
143, fig. 40 (shell). fig- 58 (radula).

Source : MNHN, Paris

190

WALTER O. CERNOHORSKY

Figs 1-6. - 1-2. Nassarius troendleorum Cernohorsky, 1980. Lagon, stn 489, 43 m ; 27.0 mm. 3-4. Nassarius bifarius
(Baird in Brenchley, 1873). Lagon, stn 471. 42 m; 17.0 mm. 5-6. Nassarius bifarius (Baird in Brenchlcv 1873»

Lagon, stn 63, 20 m ; 20.0 mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NASSARIIDAE

191

Nassarius (Nioiha) novaezelandiae (Reeve) - Cerno-
horsky, 1978 : 81, pl. 25, fig. la only; 1984 : 86.
pl. 9. fig. 4 (lectotype of N. bifarius) and fig. 6 only
[non N assit novaezelandiae Reeve, 1854].

Material examined. - Cheslerfield Islands. Chal-
cal 1 : stn DC 54. 21°26'S. 159°00'E, 36-42 m,
25 July 1984 : 1 spm. Stn DC 55, 21°24' S,
159°00'E, 55 m, 25 July 1984 : 2 spm.

New Caledonia. Lagon :

Huon Atoll : stn 438. 37 m : 3 spm. — Stn 439, 39 m :
13 spm. — Stn 441, 37 m : 1 spm. — Stn 442 +bis,

39 m : 15 spm.

Surprise Atoll : stn 446. 36 m : 3 spm. — Stn 447.

36 m : 4 spm. — Stn 448, 30 m : 1 spm. — Stn 450,

29 m : 5 spm. Stn 451. 30 m : 1 spm. — Stn 456,

37 m : 1 spm. — Stn 465, 45 m : 1 spm. Stn 466,

42 m : I spm. — Stn 467, 41 m : 2 spm. — Stn 468,

40 m : 4 spm. — Stn 469, 39 m : 6 spm. — Stn 470,

41 m : 5 spm. — Stn 471, 42 m : 5 spm.

Northern Lagoon : stn 515, 54 m : 1 spm. Stn 517,

42 m : 3 spm. — Stn 522, 42 m : 1 spm. — Stn 528,

47 m : 2 spm. Stn 529, 50 m : 13 spm. — Stn 530,

48 m : 1 spm. Stn 531, 56 m : 5 spm. Stn 532.

56 m : 9 spm. Stn 535, 46 m : 5 spm. — Stn 536,

61 m : 1 spm. — Stn 541, 45 m : 1 spm.
Southwestern Lagoon : stn 7, 14 m : 1 spm. — Stn 10.
15 m : 2 spm. Stn 36, 20 m : 1 spm. — Stn 38,

20 m : 1 spm. Stn 40, 21 m : 1 spm. — Stn 55,

23 m : 1 spm. — Stn 63, 20 m : 1 spm. — Stn 202,

13 m : 2 spm. — Stn 219, 32 m : 2 spm. — Stn 268,

24 m : 1 spm. — Stn 291, 31 m : 1 spm. — Stn 295,

41 m : 1 spm. Stn 302, 17 m : 2 spm. — Stn 348,

45 m : 2 spm. Stn 559, 52 m : 3 spm.

Distribution. — From India to the Fiji
Islands.

Remarks. Cernohorsky (1984) treated
Nassarius bifarius as a smooth form of N.
novaezelandiae (Reeve). Prolific material of N.
bifarius from New Caledonia clearly shows that
N. bifarius is a valid species, separate from N.
novaezelandiae. N. bifarius lacks granules on the
body whorl which is smooth and distinctly or
obsoletely axially ribbed and the anterior half
has a few spiral cords ; the suturai nodules are
stronger and more distinct on the last 2 whorls,
the aperture is wider and the colour is fawn to
orange-brown with a whitish columellar callus,
in contrast to the blackish-brown and white
colour pattern of N. novaezelandiae.

Nassarius (Niotha) conoidalis
(Deshayes in Bélanger, 1832)

Buccinum conoidale Deshayes in Bclanger, 1832 : 433,
pl. 3, figs 6, 7.

Nassarius (Niotha) conoidalis - Cernohorsky, 1984 :
78, pl. 7, figs 3-10 (synonymy).

Material examined. — New Caledonia. Musor-
stom 4 : stn DW 203. 22°36' S, 167°05' E. 105-110 m.
27 September 1985 : 1 spm. — Stn DW 204. 22°37' S,
167°06'E, 120 m, 27 September 1985 : 6 spm.
Lagon :

Huon Atoll : stn 439, 39 m : 1 spm. — Stn 440bis,
39 m : 1 spm. — Stn 442 +bis, 39 m : 2 spm.
Surprise Atoll : stn 447, 36 m : 4 spm. — Stn 469.
39 m : 6 spm.

Northern Lagoon : stn 478. 35 m : 1 spm. Stn 485,

32 m : 1 spm. Stn 503. 64 m : 8 spm. Stn 503bis.

66 m : 8 spm. Stn 504, 45 m : 3 spm. — Stn 516.

48 m : 1 spm. — Stn 517, 42 m : 14 spm. — Stn 522,

42 m : 6 spm. Stn 529, 50 m : I spm. — Stn 534.

48 m : 2 spm. — Stn 535, 46 m : 45 spm. — Stn 536,

61 m : 87 spm. — Stn 537. 200 m : 8 spm. — Stn 538,
195 m : 1 spm. — Stn 541. 45 m : 6 spm.
Southwestern Lagoon : stn 69, 13 m : 3 spm. Stn 80.

33 m : 1 spm. — Stn 120. 46 m : 1 spm. Stn 149.
48 m : 10 spm. — Stn 152, 23 m : 1 spm. — Stn 187,
13 m : 2 spm. Stn 192, 18 m : 1 spm. — Stn 202,

13 m : 1 spm. — Stn 219, 32 m : 1 spm. — Stn 221, 55-
65 m : 7 spm. — Stn 230, 35 m : 3 spm. — Stn 234,
56 m : 31 spm. — Stn 234bis. 60 m : 12 spm.

Stn 235, 70 m : 3 spm. — Stn 236, 67 m : 1 spm.

Stn 240, 42 m : 8 spm. — Stn 244. 37 m : 5 spm.

Stn 245, 62 m : 2 spm. — Stn 268. 24 m : 1 spm.

Stn 281, 10 m : 1 spm. — Stn 297, 30 m : 1 spm. —

Stn 301, 46 m : 2 spm. — Stn 305, 26 m : 1 spm.

Stn 306, 38 m : 1 spm. Stn 308, 18 m: 1 spm. —

Stn 315, 50 m : 5 spm. — Stn 316. 68 m : 11 spm. —
Stn 317, 66 m : 5 spm. Stn 318, 71 m : 3 spm. —

Stn 319, 75 m : 10 spm. Stn 326, 67 m : 2 spm. —

Stn 329, 80 m : 2 spm. — Stn 332, 80 m : 15 spm. —
Stn 333, 71 m : 26 spm. — Stn 334, 47 m : 2 spm.

Stn 336, 26 m : 1 spm. — Stn 346, 40 m : 2 spm.

Stn 347, 46 m : 6 spm. Stn 348, 45 m : 13 spm.

Stn 350, 67 m : 19 spm. — Stn 352, 82 m : 26 spm.

Stn 354. 78 m : 7 spm. — Stn 356, 78 m : 1 spm. —

Stn 357, 77 m : 6 spm. - Stn 359, 74 m : 1 spm.

Stn 368, 70 m : 1 spm. — Stn 370, 127 m : 1 spm. —

Stn 376. 75-76 m : 32 spm. — Stn 384. 70 m :

1 spm. — Stn 397, 125 m : 1 spm. - Stn 400. 64 m :

4 spm. — Stn 401. 49 m : 2 spm. — Stn 403, 45 m :

4 spm. — Stn 405, 27 m : 9 spm. — Stn 406, 24 m :

2 spm. Stn 407. 24 m : 3 spm. — Stn 408. 18 m:

1 spm. — Stn 410. 35 m : 18 spm. Stn 411. 40 m :

1 spm. Stn 414. 60 m : 18 spm. Stn 559. 52 m :

6 spm. — Stn 560. 48 m : 4 spm. — Stn 562. 48 m :

1 spm. Stn 564. 35 m : 3 spm. — Stn 569. 62 m :

1 spm. — Stn 571. 40 m : 2 spm. — Stn 574, 54 m :

1 spm. — Stn 575, 62 m : 1 spm. — Stn 577, 60 m :

1 spm. Stn 580, 95-100 m : 1 spm. — Stn 581.
23 m : 5 spm. — Stn 582, 67 m : 1 spm. — Stn 591.

14 m : 4 spm. — Stn 596, 35 m : 6 spm.

Eastern Lagoon : stn 601, 47-48 m : 5 spm. Stn 603,
78-80 m : 6 spm. — Stn 604, 80 m : 11 spm. —

Stn 605, 65-70 m : 1 spm. — Stn 606. 46-48 m :

3 spm. — Stn 608, 50-56 m : 10 spm. — Stn 614, 48-
50 m : 3 spm. Stn 615, 56-60 m : 6 spm. Stn 618,

Source : MNHN, Paris

192

WALTER O. CERNOHORSKY

53-58 m : 1 spm. Stn 621, 55-56 m : 5 spin.

Stn 623, 32-40 m : 1 spm. — Stn 626, 47-48 m :
3 spm. — Stn 629, 47-48 m : 1 spm. — Stn 632, 44-
45 m : 5 spm. — Stn 633, 50 m : 10 spm. — Stn 641,
50-52 m : 4 spm. — Stn 654, 32 m : 2 spm. — Stn 658,
49-51 m : 5 spm. — Stn 660, 48-52 m : 5 spm. —
Stn 662, 50 m : 3 spm. — Stn 667, 33-37 m :
29 spm. — Stn 668, 40 m : 2 spm. — Stn 669, 30-
40 m : 1 spm. Stn 687, 37-40 m : 4 spm. — Stn 688,
36-40 m : 11 spm. — Stn 692, 44-48 m : 1 spm. —
Stn 698, 40-43 m : 4 spm. Stn 701, 36-39 m :
3 spm. — Stn 702, 37 m : 4 spm. — Stn 703, 38-40 m :

1 spm. — Stn 707. 34-38 m : 7 spm. — Stn 714, 37-
38 m : 1 spm. — Stn 716, 30 m : 3 spm. Stn 723,
45 m : II spm. Stn 724, 36-38 m : 2 spm.

Stn 729, 42-45 m : 23 spm. — Stn 730, 40-43 m :

2 spm. — Stn 731, 37-42 m : 4 spm. — Stn 736, 44-
45 m : 2 spm. - Stn 737, 49-50 m : 3 spm. Stn 738,
59-61 m : 2 spm. — Stn 741, 77-80 m : 1 spm.

Distribution. — From the Red Sea and the
Persian Gulf to Fiji and the Line Islands.

Remarks. — The species was previously
known to occur subtidally to a depth of 120 m.
This range is now extended to 200 m.

A ’assarius (Niotha) délicat us (A. Adams, 1852)

Nassa delicata A. Adams, 1852 ; 99.

Nassa nodicosiata A. Adams, 1852 : 99.

Nassarius (Niotha) delicatus - Cernohorsky, 1984 :
100, pl. 14. figs 4-8.

Material examined. New Caledonia. Lagon :
Huon Atoll : stn 438, 37 m : 38 spm. Stn 439, 39 m :
1 spm. — Stn 440bis, 39 m : 17 spm. Stn 441,
37 m : 12 spm. — Stn 442 +bis, 39 m : 24 spm. —
Stn 443, 40 m : 5 spm.

Surprise Atoll : stn 446, 36 m : 36 spm. — Stn 447,
36 m : 1 spm. Stn 448, 30 m : 19 spm. — Stn 449,
21 m : 6 spm. — Stn 450, 29 m : 2 spm. — Stn 452,

27 m : 20 spm. — Stn 453, 26 m : 19 spm. —

Stn 454, 36 m : 7 spm. — Stn 455, 40 m : 7 spm.

Stn 456, 37 m : 2 spm. Stn 463, 43 m : 3 spm. —

Stn 465, 45 m : 11 spm. — Stn 466, 42 m : 3 spm.
Stn 467, 41 m : 1 spm. — Stn 469, 39 m : 18 spm. —
Stn 470, 41 m : 8 spm. — Stn 472. 48 m : 1 spm.

Stn 473, 50 m : 11 spm. Stn 474, 52 m : 1 spm.
Northern Lagoon : stn 478, 35 m : 32 spm. Stn 481,
33 m : 49 spm. Stn 482, 33 m : 24 spm. —
Stn 517, 42 m : 2 spm. — Stn 519, 39 m : 10 spm.

Stn 535, 46 m : 1 spm. Stn 536, 61 m : 2 spm. —
Stn 541, 45 m : 2 spm. — Stn 542, 50 m : 3 spm.
Southwestern Lagoon : stn 7, 14 m : 32 spm. — Stn 8,
15 m : 11 spm. - Stn 9, 10 m : 1 spm. Stn 39,

19 m : 1 spm. — Stn 40, 21 m : 14 spm. — Stn 41, 28-

46 m : 2 spm. — Stn 46, 25 m : 2 spm. Stn 47,

28 m : I spm. — Stn 50, 12 m : 9 spm. — Stn 51,

10 m : 15 spm. — Stn 64, 15 m: 16 spm. — Stn 65,

24 m : 39 spm. — Stn 66, 15 m : 13 spm. — Stn 77,

22 m : 2 spm. Stn 78, 35 m : 2 spm. — Stn 80!

33 m : 32 spm. Stn 83, 22 m : 24 spm. — Stn 84,

17 m : 5 spm. Stn 85, 21 m : 1 spm. Stn 95,

14 m : 8 spm. — Stn 98, 15 m : 7 spm. — Stn 150, 62-
68 m : 2 spm. Stn 160, 10 m : 1 spm.

Stn 161, 20 m : 14 spm. Stn 162, 10 m :
12 spm. Stn 163, 15 m : 28 spm. — Stn 170,
22 m : 19 spm. — Stn 186, 11 m : 17 spm. —
Stn 187, 13 m : 2 spm. Stn 192, 18 m : 8 spm. —

Stn 199. 50 m : 1 spm. — Stn 200, 18 m : 5 spm. —

Stn 202, 13 m : 1 spm. Stn 206, 8 m : I spm. —
Stn 212, 10 m : 29 spm. - Stn 214, 12 m : 34 spm.
Stn 217, 16 m : 9 spm. Stn 218, 15 m : 2 spm.

Stn 219, 32 m : 2 spm. Stn 226, 28 m : 31 spm. —
Stn 233, 30 m : 5 spm. - Stn 239, 43 m : 3 spm. —

Stn 265, 15 m : 3 spm. — Stn 266, 19 m : 1 spm. —

Stn 268, 24 m : 10 spm. Stn 269, 20 m : 1 spm. —
Stn 277, 30 m : 1 spm. Stn 281, 10 m : 5 spm.

Stn 284, 6 m : 2 spm. — Stn 290, 11 m : 6 spm. —
Stn 291, 31 m : 1 spm. Stn 293, 20 m : 20 spm. —
Stn 294, 21 m : I spm. - Stn 296, 26 m : 1 spm. —

Stn 305, 26 m : 4 spm. Stn 306, 38 m : 1 spm. —

Stn 308, 18 m : 2 spm. — Stn 311, 36 m : 3 spm. —

Stn 316. 68 m : 1 spm. Stn 336, 26 m : 2 spm.

Stn 339, 26 m : 2 spm. — Stn 340, 27 m : 1 spm. —

Stn 341, 19 m : 8 spm. Stn 345, 39 m : 2 spm. —

Stn 348, 45 m : 2 spm. Stn 544, 25 m : 12 spm.

Stn 546. 33 m : 1 spm. Stn 547, 29 m : 14 spm. —
Stn 548, 32 m : 3 spm. — Stn 549, 26 m : 5 spm. —

Stn 550, 24 m : I spm. Stn 554, 27 m : 1 spm. —

Stn 556, 30 m : 2 spm. — Stn 559, 52 m : 1 spm. —

Stn 560. 48 m : 1 spm. Stn 564, 35 m : 1 spm. —

Stn 571. 40 m : 2 spm. — Stn 581, 23 m : 8 spm. —

Stn 582, 67 m : 1 spm. — Stn 585, 43 m : 1 spm.

Stn 589, 31 m : 3 spm. — Stn 590, 20 m : 14 spm. —
Stn 591, 14 m : 6 spm. Stn 593, 25 m : 1 spm.
Eastern Lagoon : stn 626, 47-48 m : 4 spm. — Stn 632,
44-45 m : 9 spm. Stn 633, 50 m : 3 spm. — Stn 641,
50-52 m : 2 spm. Stn 664. 28-30 m : 1 spm. Stn
675, 43 m : I spm. — Stn 682, 36-37 m : 4 spm —
Stn 688, 36-40 m : 2 spm. Stn 693, 35-38 m :
1 spm. — Stn 696, 41-57 m : 10 spm. Stn 697, 35-

36 m : 4 spm. Stn 698, 40-43 m : 1 spm. Stn 702,

37 m : 14 spm. - Stn 703, 38-40 m : 8 spm. — Stn

709, 39-40 m : 4 spm. Stn 713, 34-35 m : 1 spm._

Stn 715. 34-35 m : 1 spm. Stn 724, 36-38 m : 3 spm.

Stn 730. 40-43 m : 10 spm.

Distribution. From the Red Sea and the
Persian Gulf to the Ryukyu and Fiji Islands.

Remarks. This species has been dredged in
great numbers, and the majority of specimens
were the nodulose form nodicosiata A. Adams,
although a few specimens were the typical delica¬
tus form. In the nodulose form the axial ribs on
the body whorl are distinctly nodulose and form
5-6 spiral rows of nodules.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NASSAR1IDAE

193

N as sari us ( Niotha ) ecstilbus
(Melvill & Standen, 1896)

Nassa (Telasco) ecstilbus Melvill & Standen, 1896 :
274, pl. 9, fig. 4 (Type locality : Lifu, Loyalty
Islands).

Nassarius (Niotha) ecstilbus - Cernohorsky, 1984 :
101, pl. 14. figs 9, 10; pl. 15, fig. 1.

Material examined. New Caledonia. Lagon ;
Surprise Atoll : stn 449, 21 m : 5 spm. — Stn 452,
27 m : 2 spm.

Northern Lagoon : stn 481, 33 m : 4 spm. — Stn 482,
33 m ; 2 spm. — Stn 486, 33 m : 1 spm.
Southwestern Lagoon : stn 7, 14 m : 1 spm. — Stn 49,

10 m : 2 spm. Stn 50, 12 m : 1 spm. Stn 98,
15 m : 5 spm. Stn 162, 10 m : 2 spm. — Stn 186,

11 m : 4 spm. — Stn 214, 12 m : 1 spm. — Stn 221, 55-
65 m : I spm. Stn 255, 11 m : 1 spm. — Stn 283,
13 m : 2 spm. Stn 293, 20 m ; 1 spm. — Stn 308,
18 m : 3 spm.

Distribution. From the Red Sea to the
Philippines and the Fiji Islands.

Remarks. — This species has been dredged in
New Caledonian waters to a depth of 65 m.

Nassarius (Niotha) fret or um
(Melvill & Standen. 1899)

Nassa fretorum Melvill & Standen, 1899 : 159, pl. 10,
fig. ‘3.

Nassarius (Niotha) fretorum - Cernohorsky. 1984 :
101, pl. 15. figs 2-5.

Material examined. New Caledonia. Lagon ;
Southwestern Lagoon : stn 324, 39 m : 1 spm.

Stn 358, 50 m : 1 spm.

Eastern Lagoon : stn 597 bis, 50-70 m : 1 spm.

Distribution. From India to the Philip¬
pines and NE Australia ; now New Caledonia.

Remarks. — This is a new geographical
record for the species.

Nassarius ( Niotha ) himeroessa
(Melvill & Standen, 1903)

Nassa (Alectrvon) himeroessa Melvill & Standen,
1903 : 306, pl. 22, fig. 7.

Nassarius (Niotha) himeroessa - CERNOHORSKY, 1984 :
107, pl. 17, figs 5-7.

Material examined. Chesterfield Islands. Mus-
ORSTOM 5 : stn 300. 22°48' S. I59"24' E. 450 m.

11 October 1986 : 1 spm. — Stn 304, 22° 10' S,
159°26' E, 385-420 m, 12 October 1986 : 1 spm. Stn
306, 22°08' S. 159°21 ' E. 375-415 m. 12 October 1986 :

1 spm.

New Caledonia. Biocal : stn DW 08, 20“34' S,
166°54' E. 435 m, 12 August 1985 : 5 spm. — Stn DW
33, 23" 10' S, 167° 10' E, 29 August 1985 : 2 spm. - Stn
DW 46, 22°53'S, 167°17'E. 570-610 m, 30 August
1985 ; 1 spm. Stn DW 48. 23W S. 167"29' E. 775 m.
31 August 1985 : 9 spm. - Stn DW 49. 23°03' S.
167"32' E. 824-830 m, 31 August 1985 ; 1 spm. Stn
DW 51, 23°05'S. I67°45’E, 680-700 m. 31 August
1985 : 23 spm. Stn DW 56, 23°35' S. 167" 12' E. 695-
705 m, 1 September 1985 ; 1 spm. Stn DW 66. 24“
55' S, 168°22' E, 505-515 m, 3 September 1985 : 8 spm.

— Stn DW 77, 22° 15' S, 167° 15' E, 440 m. 5 Septem¬
ber 1985 : 22 spm. Stn DW 83, 20°35' S. 166’54’ E.
460 m, 6 September 1985 : 1 spm. — Stn DW 106, 21°
36' S, 166"29' E, 625-650 m, 8 September 1985 ; 1 spm.

Musorstom 4 : stn DW 149. 19°08'S. 163°23'E,
155 m, 14 September 1985 : 1 spm. — Stn DW 156.
I8°54' S. 163° 19' E, 525 m. 15 September 1985 : 1 spm.

— Stn DW 159, 18"46'S. 163°16'E, 585 m. 15
September 1985 : 1 spm. — Stn CC 247, 22°09'S,
167° 13' E, 435-460 m. 4 October 1985 ; 1 spm.

Distribution. — From the Persian Gulf to
the Philippines and the Kermadec Islands.

Remarks. — This is a new record of the
species for New Caledonia.

Nassarius (Niotha) quadrasi
(Hidalgo, 1904)

Nassa gruneri — Reeve, 1853 : pl. 12, fig. 75 (non
Buccinum gruneri Dunker. 1846).

Nassa quadrasi Hidalgo, 1904 : 204 (nom. nov. pro
Nassa gruneri of Reeve, 1853).

Nassarius (Niotha) quadrasi - Cernohorsky, 1984 :
108. pl. 17. figs 8-9 (synonymy).

Material examined. New Caledonia. Lagon :
Surprise Atoll : stn 453, 26 m ; 2 spm.
Southwestern Lagoon : stn 9. 10 m : 1 spm. — Stn 162,
10 m : 4 spm. Stn 214, 12 m : 4 spm. Stn 226.

28 m : 3 spm. — Stn 308, 18 m ; 2 spm. Stn 341,

19 m : 1 spm. Stn 591, 14 m : 1 spm.

Distribution. — From Malaysia to the Ryu-
kyu and Samoa Islands.

Nassarius (Niotha) sinusigerus
(A. Adams, 1852)

Nassa sinusigera A. Adams, 1852 : 100.

Nassarius (Niotha) sinusigerus - Cernohorsky, 1984 :
103, pl. 16, figs 1-10 (synonymy).

Source : MNHN, Paris

194

WALTER O. CERNOHORSKY

Material examined. — New Caledonia. Lagon :
Southwestern Lagoon : stn 34, 10 m : 1 spm.

Stn 139, 45 m : 6 spm. — Stn 140, 47 m : 4 spm.

Stn 141. 44 m : 19 spm. Stn 142, 34 m : 6 spm.
Stn 143, 32 m : 23 spm. — Stn 144, 25 m : 21 spm.
Stn 278, 17 m : 1 spm.

Eastern Lagoon : stn 721, 22-23 m : 1 spm.

Distribution. Front the Red Sea and the
Gulf of Oman to the Fiji Islands.

Remarks. — This is a new record of the
species for New Caledonia.

N’assarius (Niotha) splendidulus
(Dunker, 1846)

Fig. 7

Buccinum splendidulum Dunker, 1846 : 170.
Nassarius (Niotha) splendidulus - Cernohorsky,
1984 : 85. pl. 8, figs 8-9 and 11-13 ; pl. 9, figs 1-2
(synonymy — with the exclusion of references to
Nassarius stigmarius A. Adams. 1852).

Material examined. — New Caledonia. Lagon :
Huon Atoll : stn 438, 37 m : 2 spm. — Stn 440 bis,
39 m : 1 spm. — Stn 441, 37 m : 4 spm.

Stn 442+ bis, 39 m : 1 spm. — Stn 443. 40 m : 1 spm.
Surprise Atoll : stn 446, 36 m : 5 spm. — Stn 448,
30 m : 2 spm. — Stn 449, 21 m : 7 spm. Stn 452,

27 m : I spm. — Stn 453, 26 m : 2 spm. — Stn 454,

36 m : 2 spm. Stn 455, 40 m : 5 spm. Stn 463,

43 m : 2 spm. — Stn 465, 45 m : 2 spm. — Stn 469,

20 m : 2 spm. Stn 470, 41 m : 2 spm. — Stn 472,

48 m : 1 spm. Stn 473, 50 m : 1 spm.
Northern Lagoon : stn 478, 35 m : 3 spm. Stn 481,
33 m : 8 spm. Stn 482, 33 m : 8 spm. — Stn 484,

35 m : 1 spm. — Stn 485, 32 m : 2 spm. — Stn 504,

45 m : 4 spm. Stn 517, 42 m : 2 spm. — Stn 522,

42 m : 2 spm. Stn 533, 50 m : 1 spm. Stn 536,

61 m : 1 spm. Stn 541. 45 m : 2 spm. - Stn 542,

50 m : 7 spm.

Southwestern Lagoon : stn 7, 14 m : 3 spm. Stn 40,

21 m : 10 spm. — Stn 51. 10 m : 2 spm. Stn 66.
15 m : 2 spm. Stn 80, 33 m : 5 spm. — Stn 83,

22

m

10 spm.

Stn 84, 17 m

: 4 spm.

Stn 95,

14

m

2 spm.

— Stn 98, 15 m

4 spm.

Stn

101,

18

m

1 spm.

Stn 120, 46 m

1 spm.

Stn

121,

12

m

3 spm. —

Stn 150, 62-68 m

: 6 spm. -

Stn

152,

23

m

1 spm.

Stn 161. 20 m

2 spm.

Stn

163,

15

m

4 spm.

- Stn 170. 22 m

7 spm. —

Stn

192,

18

m

1 spm.

Stn 199, 50 m

1 spm. -

Stn

200,

18

m

5 spm.

Stn 201, 17 m :

48 spm. -

Stn

202,

13

m

1 spm.

- Stn 211, 12 m

1 spm.

Stn

214.

12

m

1 spm.

Stn 217. 16 m

3 spm.

Stn

218,

15

m

12 spm.

- Stn 226, 28 m

6 spm. —

Stn

233,

30

m

4 spm.

Stn 239, 43 m

2 spm.

Stn

241,

35

m

1 spm.

- Stn 249, 11 m

1 spm. —

Stn

253.

16

m

1 spm.

Stn 268, 24 m

5 spm.

Stn

275,

19 m : 1 spm. — Stn 281, 10 m : I spm. Stn 284,
6 m : 1 spm. Stn 291, 31 m : 4 spm. Stn 293^

20 m : 8 spm. Stn 294, 21 m : I spm. Stn 300,

21 m : I spm. Stn 304, 27 m : 4 spm. Stn 305,

26 m : 3 spm. — Stn 311, 36 m : I spm. Stn 313,

30 m : 1 spm. — Stn 324, 39 m : 5 spm. — Stn 334*

47 m : 2 spm. Stn 336, 26 m : 8 spm. — Stn 338,

32 m : I spm. Stn 339, 26 m : 2 spm. — Stn 340,

27 m : 3 spm. Stn 341, 19 m : I spm. - Stn 344,

37 m : 2 spm. Stn 345, 39 m : 1 spm. Stn 348^

45 m : 4 spm. Stn 405, 27 m : 2 spm. — Stn 410,

35 m : 10 spm. Stn 544, 25 m : 10 spm. Stn 546,

33 m : 2 spm. Stn 547, 29 m : 2 spm. Stn 548,

32 m : I spm. Stn 555, 32 m : 2 spm. Stn 556,

30 m : 1 spm. Stn 581, 23 m : 3 spm. — Stn 589,

31 m : 2 spm. Stn 590, 20 m : 6 spm. - Stn 591,

14 m : 4 spm. Stn 593, 25 m : I spm.

Eastern Lagoon : stn 620, 50-52 m : I spm. — Stn 625,
34-40 m : 1 spm. Stn 631, 43 m : 3 spm. Stn 632,

44-45 m : 5 spm. Stn 633, 50 m : 2 spm. Stn 641,

50-52 m : 3 spm. Stn 681, 33 m : 3 spm. Stn 682’

36-37 m : 5 spm. Stn 685, 24-26 m : 1 spm. —
Stn 697, 35-36 m : 4 spm. - Stn 701, 36-39 m :
1 spm. Stn 702, 37 m : 12 spm. Stn 703, 38-
40 m : 3 spm. Stn 709, 39-40 m : 2 spm. Stn 713,
34-35 m : I spm. Stn 715, 34-35 m : 1 spm. —
Stn 716, 30 m : 2 spm. Stn 724, 36-38 m : 1 spm. —
Stn 729,42-45 m : 4 spm. Stn 730, 40-43 m : 2 spm.

Distribution. From the Red Sea and the
Persian Gulf to Hawaii and the Society Islands.

Nassarius (Niotha) stigmarius

(A. Adams, 1852)

Figs 8-9

Nassa stigmaria A. Adams, 1852 : 96 (Type locality :
Siquijor Id., Philippines).

Nassarius (Niotha) splendidulus - Cernohorsky [parsl
1984 : 85, pl. 8, fig. 10 only (illustrated holotype of
Nassa stigmaria A. Adams) [non Buccinum splendi¬
dulum Dunker. 1846].

Material examined. Cheslerfield Islands. Chal-
CAL 1 : stn DC 9. 20"44' S, 161“02‘ E, 75 m. 15 July
1984 : 2 spm. Stn DC 10, 20”36' S. 161°06' E, 87 m.
15 July 1984 : 2 spm. Stn DC 17, 19"12'S, 158“
56'E, 44 m, July 1984 : Il spm. Stn DC 26
19”! 1'S, 158 35' E. 48 m. 18 July 1984 : 1 spm. Stn
DC 39, 20"29' S, 158“4I' E, 40 m, 23 July 1984 : 1 spm
Stn DC 40. 20°32' S, 158"51' E, 65 m. 23 July 1984 !
2 spm. — Stn DC 41. 20“35'S, 158°47'E, 67 m, 23
July 1984 : 3 spm. Stn DC 45, 20°49' S. 158"36' E
50 m, 23 July 1984 : 2 spm. Stn DC 46. 20°52' s’
158“34' E, 65 m, 23 July 1984 : 2 spm. Stn DC 49
20"58' S, 158°35' E, 48 m. 24 July 1984 : 1 spm. — Stn
DC 52, 21-13' S, 158 49' E, 69 m. 24 July 1984 : 1 spm
Stn DC 55, 21-24' S. 159“00' E, 55 m. 25 July 1984 :
4 spm. Stn DC 56. 21’24' S. I59°09' E, 60 m, 25
July 1984 : 3 spm. Stn DC 59, 21"40' S. 159“21' E,

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NASSARUDAE

195

Figs 7-12. 7. Nassarius splendidulus (Dunker. 1846). Lagon, stn 83. 22 m; 14.0 mm. 8-9. Nassarius sligmarius

(A. Adams, 1852). Lagon, stn 542, 50 m ; 15.0 mm. 10. - Nassarius vidaknsis (Barnard. 1959). Biocal. stn DW 64.
250 m ; 7.3 mm. 11. Nassarius abyssicolus (A. Adams, 1852). Chalcal 1. stn DC 26. 48 m ; 11.0 mm. 12.
Nassarius haldemanni (Dunker, 1847). Lagon, stn 620, 50-52 m; 16.0 mm.

56 m, 25 Julv 1984 : 1 spm. Stn CP 16, 21°42' S.
159°22' E, 5.3 m, 25 July 1984 : I spm. — Stn DC 61,
21"42'S, 159"29'E, 50 m. 26 July 1984 : 5 spm.

New Caledonia. Lagon :

Huon Atoll : stn 433, 40-67 m : 1 spm.

Surprise Atoll : stn 446, 36 m : 1 spm. — Stn 455,
40 m : 1 spm.

Northern Lagoon : stn 542, 50 m : 2 spm.
Southwestern Lagoon : stn 200, 18 m : 1 spm. — Stn
229, 41 m : 1 spm. — Stn 264, 56 m : 1 spm.

Stn 267, 70 m : 2 spm. - Stn 310, 46 m : 1 spm.

Stn 340, 27 m : 1 spm. Stn 546, 33 m : 1 spm.
Stn 570, 53 m : 1 spm. — Stn 572. 65 m : 1 spm.
Eastern Lagoon : stn 598, 73-75 m : 2 spm. — Stn 644,
45-48 m : 1 spm. — Stn 736, 44-45 m : 1 spm.

Distribution. From the Philippines to
Japan and New Caledonia. subtidal, from 18-
87 m.

Source : MNHN, Paris

196

WALTER O. CERNOHORSKY

Remarks. Cernohorsky (1984) erroneously
placed N. (N.) stigmarius (A. Adams), in syno-
nymy of N. (N.) splendidulus (Dunker). The
large number of specimens of both species taken
in New Caledonian waters clearly shows that
they are separate species. N. (N.) stigmarius is
lighter in weight and considerably less solid than
N. (N.) splendidulus , with a finer sculpture
consisting of close-set nodules, thinner outer lip
and a smaller pariétal callus which is thinning
posteriorly ; it is cream in colour with small
undefined brown spots at sutures, 3 brown larval
whorls and only 1 post-larval whorl purplish-
brown. N. (N.) splendidulus always lacks the
purplish-brown colouring of the apical whorls.

Nassarius (Telasco) multipunctatus
(Schepman, 1911)

Nussa (Zeuxis) multipunctata Schepman, 1911 : 321,
pl. 20, figs 4a. b.

Nassarius (Telasco) multipunctatus - Cernohorsky,
1984 : 122, pl. 22, figs 1-2 (synonymy).

Material examined. — New Caledonia. Musor-
stom 4 : stn DW 187, 19°08' S, 163“29' E, 65-120 m,
19 September 1985 : I spm. - Stn CP 190, 19°06' S,
163“29' E, 215 m. 19 September 1985 : 3 spm.

Stn DW 227, 22°46' S, 167“20' E. 300 m, 30 September
1985 : 1 spm.

Lagon :

Huon Atoll : stn 433, 40-67 m : 1 spm.

Northern Lagoon : stn 495, 80 m : 1 spm. — Stn 536,
61 m : 3 spm. — Stn 537, 200 m : 1 spm. — Stn 542,
50 m : 5 spm.

Southwestern Lagoon : stn 234, 56 m : 2 spm. —
Stn 234 bis, 60 m : 2 spm. — Stn 239, 43 m : 2 spm. —
Stn 240, 42 m : 6 spm. — Stn 244, 37 m : 1 spm. —

Stn 319. 50 m : 6 spm. — Stn 320, 70 m : 3 spm.

Stn 322, 71 m : 2 spm. Stn 324; 39 m : 8 spm.

Stn 331, 79 m : 11 spm. — Stn 333, 71 m : 1 spm.

Stn 334. 47 m : 1 spm. Stn 344, 37 m : 1 spm.

Stn 348. 45 m : 3 spm. — Stn 350, 67 m : 1 spm.

Stn 353, 70 m : 1 spm. Stn 357, 77 m : 4 spm.

Stn 358, 50 m : I spm. Stn 360. 60 m : 1 spm. —
Stn 361, 78 m : 6 spm. — Stn 363, 67 m : 2 spm. —
Stn 368, 70 m : 3 spm. — Stn 374, 70-72 m : 2 spm. —
Stn 375, 67-71 m : 2 spm. Stn 376, 75-76 m :
1 spm. Stn 377, 56 m : 2 spm. — Stn 383, 62 m :
1 spm. — Stn 384, 70 m : 2 spm. — Stn 385, 75 m :
3 spm. Stn 391, 65 m . 1 spm. Stn 392, 80 m :
1 spm. — Stn 398, 71 m : 4 spm. — Stn 400. 64 m :
3 spm. — Stn 403, 45 m : 12 spm. — Stn 405, 27 m :
I spm. — Stn 413. 40-60 m : 3 spm. Stn 414, 60 m :
7 spm. — Stn 428, 56 m : 3 spm. Stn 429, 95 m :

1 spm. — Stn 561. 48 m : spm. — Stn 562, 48 m :

2 spm. — Stn 580, 95-100 m : 2 spm. — Stn 589,
31 m : 2 spm.

Eastern Lagoon : stn 598, 73-75 m : 20 spm. —

Stn 599, 50 m : I spm. Stn 601, 47-48 m : 1 spm._

Stn 602, 43-48 m : 5 spm. — Stn 604, 80 m : 1 spm. —

Stn 607, 48-54 m : 7 spm. Stn 615, 56-60 m :
1 spm. Stn 620. 50-52 m : 1 spm. — Stn 621, 55-
56 m : 2 spm. Stn 626, 47-48 m ; 1 spm. Stn 633,

50 m : 1 spm. Stn 640. 50-80 m : 4 spm. — Stn 662^

50 m : 1 spm. — Stn 668, 40 m : 1 spm. — Stn 729, 42-
45 m : 2 spm. — Stn 736, 44-45 m : 1 spm. — Stn 737,
49-50 m : 1 spm.

Distribution. From East Africa to the
Philippines and the Solomon Islands ; now New
Caledonia.

Remarks. The record of N. (T.) multipunc¬
tatus from New Caledonia is a new eastward
range extension. The bathymétrie range is ex-
tended from 200 m to 300 m.

Nassarius (Telasco) shacklefordi
(Melvill & Standen, 1896)

Nassa (Telasco) shacklefordi Melvill & Standen,
1896 : 274, pl. 9, fig. 3 (Type locality : Lifou,
Loyalty Islands).

Nassarius (Telasco) shacklefordi - Cernohorsky,
1984 : 123, pl. 22, figs 6-8 (synonymy).

Material examined. Chesterfield Islands. Chal-
cal 1 : stn DC 6, 20°57' S, 161"43' E, 45 m. 14 July
1984 : I spm. Stn DC 7, 20“51' S, 161°37' E, 62 m,
14 July 1984 : 1 spm. Stn DC 9, 20°44' S, 16T02' e!
75 m, 15 July 1984 : 1 spm. Stn DC 17. 19°12'S
158“56'E, 44 m, 17 July 1984 : 3 spm. Stn DC 24
19° 11 ' S, 158-37'E, 38 m, 18 July 1984 : 6 spm —
Stn DC 25. 19-09'S, 158"32' E. 56 m, 18 July 1984 •
1 spm. Stn DC 45, 20°49' S, 158°30' E, 50 m, 23
July 1984 : 4 spm. Stn DC 55, 21 "24' S, 159"00' E,

55 m, 25 July 1984 : 1 spm. Stn DC 60, 21°49' S
159“28' E, 45 m, 25 July 1984 : 3 spm.

New Caledonia. Lagon :

Southwestern Lagoon : stn 7, 14 m : 1 spm. — Stn 9,
10 m : 1 spm. Stn 64, 15 m: 1 spm. Stn 79,

16 m : 6 spm. — Stn 80, 33 m : 1 spm. — Stn 82,

10 m : 1 spm. — Stn 83, 22 m : 3 spm. Stn 84,

17 m : 1 spm. Stn 95. 14 m : I spm. Stn 99,

14 m : 3 spm. Stn 192, 18 m : 1 spm. — Stn 217,

16 m : 2 spm. Stn 226, 28 m : 1 spm. Stn 264*

56 m : 2 spm. - Stn 267, 70 m : 3 spm. — Stn 293^

20 m : 5 spm. — Stn 294, 21 m : 4 spm. Stn 296^

26 m : 3 spm. Stn 324, 39 m : 2 spm. — Stn 342,

55 m : 1 spm. - Stn 343, 32 m : 1 spm. — Stn 545,

37 m : 7 spm. Stn 550, 24 m : I spm. — Stn 552^

38 m : 1 spm. — Stn 561, 48 m : 1 spm.

Eastern Lagoon : stn 597 bis, 50-70 ni : I spm. _

Stn 677, 32 m : 1 spm

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NASSAR1IDAE

197

Distribution. From Lord Howe Island,
Australia to New Caledonia and tentatively
Japan.

Remarks. — The known bathymétrie range
for the species is extended from 40 m to 75 m.

Nassarius (Telasco) vidalensis (Barnard, 1959)
Fig. 10

Nassa vidalensis Barnard, 1959 : 118, textfig. 24b.
Nassarius (Telasco) vidalensis - Cernohorsky, 1984 :
123, pl. 22, figs 3-5; 1988 : 84.

Material examined. Chesterfield Islands. Chal-
CAL 1 ; stn DC 53, 21°19' S, 158*55' E, 60 m, 24 July
1984 ; 1 spm.

Musorstom 5 ; stn 298, 22"44' S, 159°22' E, 320 m,
11 October 1986 : 1 spm. Stn 299, 22°48' S,
159 u 24' E, 360-390 m, 11 October 1986 : 1 spm.

Stn 302, 22° 10' S, 159°23'E, 345-360 m. 12 October
1986 ; 1 spm. — Stn 304, 22*10'S, 159*26'E, 385-
420 m, 12 October 1986 ; 4 spm.

New Caledonia. Biocal : stn DW 64, 24°48' S,
168°09’E, 250 m, 3 September 1985 ; 11 spm.

Chalcal 2 : stn DW 71, 24"42' S, 168° 10' E, 230 m,
27 October 1986 : 2 spm. Stn DW 79. 23*41'S,
168°00' E, 243 m, 30 October 1986 : 1 spm. — Stn DW
84. 23°24'S, 168°07' E, 170 m, 31 October 1986 :
1 spm.

Distribution. — From Reunion Island to
South Africa, NW Australia and Japan ; now
New Caledonia.

Remarks. The record of the species from
New Caledonia represents a southeastward range
extension. The known bathymétrie range is
extended from 225 m to 420 m. Fresh specimens
are brightly ornamented with reddish-brown.

Nassarius (Zeuxis) ahyssicolus (A. Adams. 1852)
Fig. 11

Nassa abyssicola A. Adams. 1852 : 100.

Nassarius (Zeuxis) ahyssicolus - Cernohorsky, 1984 :
166, pl. 35, figs 1-3.

Material examined. Chesterfield Islands. Chal¬
cal 1 ; stn DC 26, 19° 11 ' S. 158°35' E, 48 m, 18 July
1984 : 1 spm.

Distribution. To date known only from
the Philippines and the Hawaiian Islands ; now
New Caledonia.

Remarks. — This is a new record for New
Caledonia and a Southern range extension.

Nassarius (Zeuxis) arcus sp. nov.

Figs 13-16

Type material. - Holotype in mnhn : para-

types in MNHN. AMS, AIM, NMNZ, NSMT, NM, USNM.

Type locality. — New Caledonia. Lagon :
Southweslem Lagoon, Grand Récif Sud : stn 397.
22*39'S, 167*1 l'E, 125 m, 23 January 1985.
B. Richer-orstom coll.

Material examined (ail paratypes). New Caledo¬
nia. Musorstom 4 : stn DW 149. 19°08'S, 163°23' E,
155 m, 14 September 1985 : 150 spm, Bouchet &
Richer coll. Stn DW 150. 19°07' S, 163°22' E,
110 m, 14 September 1985 : 102 spm. Bouchet &
Richer coll. Stn DW 151. 19*07'S. 163*22'E,
200 m, 14 September 1985 : 38 spm.

Lagon ; Southwestern Lagoon : stn 429, 22*40' S,
167*15' E, 95 m. 25 January 1985 : 7 spm, B. Richer-
orstom coll.

Description. Shell small, up to 9.0 mm
in length, elongate-ovate, width 55-60 % of shell
length, light in weight, teleoconch of 4-4 1/4
convex whorls, protoconch of 3 1/4-3 1/2 smooth,
glassy-white larval whorls, ultimate turn finely
carinate ; first 3 post-larval whorls angulate
centrally, angulation usually only faintly indi-
cated on the presutural ramp of the body whorl.
Sculptured with distinct axial ribs which con¬
tinue to the back of the outer lip and number
from 13-20 on the penultimate and from 14-24
on the body whorl ; spiral sculpture distinct,
consisting of fine spiral grooves which number
from 4-7 on the penultimate and from 8-12 on
the body whorl ; followed by 2-5 raised spiral
threads anteriorly and from 5-7 oblique cords on
the siphonal fasciole ; sutures with a uniformly
nodulose girdle. Aperture about the same height
as the spire, height 45-51 % of shell length. outer
lip thickened and broadly varieed, interior of
outer lip with 8-11 fine, elongated denticles,
columella concave and with 2-4 minute denticles,
plus a weak or distinct pariétal denticle, columel-
lar callus narrow and confined to aperture,
siphonal canal short, broad and prominent.
White in colour, spire whorls with 2 brownish
suturai bands. body whorl with a suturai, central

Source : MNHN, Paris

198

WALTER O. CERNOHORSKY

Figs 13-18. 13-15. — Nassarius arcus sp. nov. Holotype. Lagon, stn 397, 125 m ; 7.9 mm (15, coalcd to show sculpture).

16. — Nassarius arcus sp. nov. Paratypc. Lagon, stn 429, 95 m ; 8.4 mm (slendcr individual). 17. Cyllene concinna
A. Adams. 1851. Lagon, stn 218, 15 m : 12.5 mm. 18. Cyllene fuscata A. Adams, 1851. Musorstom 4. stn DW 150

110 m; 15.5 mm.

and sometimes basal brown band, aperture and
varix white. Some specimens are white and hâve
a single orange-brown band on body whorl.
Preserved animal white, operculum translucent
pale yellow with a short brown line in centre,
rounded and simple at margins.

Distribution. — To date known only from
New Caledonia, in 95-200 m.

Remarks. N. (Z.) arcus is only superficially
similar to N. (Z.) agapetus (Watson, 1882), but
this species lacks the features of compressed and
centrally angulate whorls and the prominent
presutural groove and suturai nodules of N. (Z.)
arcus. Some specimens of N. (Z.) arcus, espe-
cially those from Musorstom 4, stn DW 149, are
rather compressed so that ail spire whorls are
distinctly angulate on the presutural ramp and
the body whorl is also subangulate.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NASSAR1IDAE

199

Nassarius (Zeuxis) barsdelli Ladd, 1976

Nassarius (Alectrion) barsdelli Ladd, 1976 : 131,
figs 12-15.

Nassarius (Zeuxis) barsdelli - Cernohorsky, 1984 :
140. pl. 27. figs 1-3.

Material examined. New Caledonia. Lagon :
Southwestern Lagoon : stn 329, 80 m : 1 spm
immature.

Distribution. Previously known only from
Recent specimens in the Philippines and fossil
specimens from the Pleistocene of Vanuatu ; now
New Caledonia.

Remarks. Only one immature specimen of
N. (Z.) barsdelli has been taken in New Caledo-
nian waters. This is only the second locality after
the Philippines from where Recent specimens
hâve been recorded.

Nassarius (Zeuxis) castus (Gould, 1850)

Nassa casta Gould, 1850 : 154.

Nassarius (Zeuxis) castus - Cernohorsky, 1984 : 130.
pl. 24, figs 7-17; pl. 25, figs 1-10 (synonymy).

Material examined. New Caledonia. Lagon :
Northern Lagoon : stn 513, 55 m : 14 spm.
Southwestern Lagoon : stn 17, 24 m : 1 spm. — Stn 19,
29 m : 1 spm. — Stn 41. 28-46 m : 1 spm. — Stn 69,

13 m : 1 spm. Stn 74, 31 m : 1 spm. Stn 86.

29 m : 5 spm. — Stn 91, 30 m . 1 spm. — Stn 107,

33 m : 1 spm. Stn 108, 17 m : 1 spm. Stn 130,

32 m : 1 spm. Stn 131, 38 m : 3 spm. Stn 133, 59-
62 m : I spm. Stn 139. 45 m : 6 spm. — Stn 149,
48 m : 7 spm. — Stn 173, 20-50 m : 2 spm. — Stn 178,
20 m : 1 spm. Stn 184, 13 m : 1 spm. — Stn 187,

13 m : 6 spm. Stn 192, 18 m : 3 spm. — Stn 193,

20 m : 1 spm. — Stn 202, 13 m : 1 spm. — Stn 203,

13 m : 1 spm. Stn 208, 9 m : 2 spm. — Stn 209.

14 m : 6 spm. — Stn 210. 14 m : 1 spm. Stn 211,

12 m : 2 spm. Stn 215, 14 m : 14 spm. — Stn 216,
14 m : 10 spm. — Stn 219, 32 m : 9 spm. — Stn 220,
12 m : I spm. — Stn 221, 55-65 m : 15 spm. —
Stn 235, 70 m : 1 spm. — Stn 238, 50 m : 1 spm.

Stn 245, 62 m : 1 spm. — Stn 268, 24 m : 1 spm.

Stn 277, 30 m : 1 spm. — Stn 278, 17 m : 2 spm.

Stn 317, 66 m : 1 spm. Stn 318, 71 m : I spm. —

Stn 320, 70 m : 3 spm. — Stn 329, 80 m : 1 spm. —

Stn 354, 78 m : 2 spm. — Stn 550, 24 m : 4 spm. —

Stn 591, 14 m : 1 spm.

Eastern Lagoon : stn 606, 46-48 m : 2 spm. — Stn 608,
50-56 m : 9 spm. — Stn 611, 56-57 m : 3 spm. —
Stn 614, 48-50 m : 2 spm. — Stn 619, 27-42 m :
1 spm. Stn 624, 44-46 m : I spm. Stn 628. 55-
56 m : 15 spm. Stn 635, 45-52 m : 1 spm. —

Stn 636, 34-40 m : 82 spm. — Stn 638. 56-58 m :
1 spm. Stn 643, 56-66 m : 8 spm. — Stn 646, 66-
70 m : 1 spm. — Stn 647. 50-52 m : 1 spm. Stn 649,
64-65 m : 3 spm. — Stn 652, 55-62 m : 2 spm.

Stn 654, 32 m : 3 spm. Stn 655. 35-40 m : 2 spm.

Stn 656. 30-40 m : 14 spm. - Stn 660. 48-52 m :
5 spm. — Stn 667, 33-37 m : 3 spm. — Stn 670, 48 m :
1 spm. — Stn 672, 15-20 m : 2 spm. Stn 674, 48 m :
5 spm. Stn 683, 42-45 m : I spm. Stn 692, 44-
48 m : 6 spm. — Stn 694. 45-47 m : 7 spm. — Stn 695,
54-55 m : 16 spm. — Stn 705, 46-48 m : 1 spm. —
Stn 712,47-49 m : 7 spm. Stn 723, 45 m : 4 spm.

Stn 727, 45-46 m : 2 spm. — Stn 728, 43-47 m :
13 spm. — Stn 729. 42-45 m : 3 spm. Stn 732, 43-
50 m : 3 spm. — Stn 733. 35-38 m : 1 spm. — Stn 737,
49-50 m : 3 spm. — Stn 738, 59-61 m : 14 spm.

Distribution. - From the Red Sea and the
Persian Gulf to Japan and the Samoa Islands.

Remarks. The majority of the New Caledo-
nian N. (Z.) castus populations belong to the
form vitiensis Rousseau, 1854.

Nassarius (Zeuxis) complus (A. Adams, 1852)

Nassa compta A. Adams, 1852 : 107.

Nassarius (Zeuxis) complus - Cernohorsky : 146.
pl. 29. figs 1-7 (synonymy).

Material examined. Chesterfield Islands. Ciial-
cal 1 : stn DC 44. 20°46' S. 158°34' E. 79 m. 23 July
1984 : 7 spm.

New Caledonia. Musorstom 4 : stn CP 148.
19°23' S, 163°32' E, 58 m, 14 September 1985 : 2 spm.
Lagon :

Northern Lagoon : stn 506, 56 m : 1 spm. — Stn 529,
50 m : 5 spm. Stn 532, 56 m : 2 spm. Stn 535,
46 m : 2 spm. — Stn 539, 240 m : 2 spm.
Southwestern Lagoon : stn 1. 19 m : 3 spm. — Stn 2.
14 m : 2 spm. Stn 11, 24 m : 16 spm. Stn 16.

30 m : 3 spm. — Stn 17. 24 m : 3 spm. — Stn 19,

29 m : 4 spm. Stn 24. 28 m : 4 spm. Stn 25,

28 m : 13 spm. Stn 26, 22 m : 1 spm. — Stn 27,

18 m : 3 spm. Stn 28, 9 m : 1 spm. Stn 29, 12 m :

6 spm. — Stn 31, 29 m : 2 spm. — Stn 32, 30 m :

5 spm. — Stn 33, 18 m : 4 spm. — Stn 34, 10 m :

5 spm. — Stn 41, 28-46 m : 2 spm. Stn 42. 25 m :
3 spm. — Stn 45, 14 Yn : 2 spm. Stn 46, 25 m :

3 spm. — Stn 47, 28 m : 2 spm. — Stn 54, 25 m :

5 spm. Stn 55, 23 m : I spm. Stn 57, 10 m :

2 spm. Stn 58, 22 m : 1 spm. — Stn 69. 13 m :

1 spm. — Stn 70, 30 m : 1 spm. Stn 71, 22 m :

5 spm. Stn 72. 15 m : 6 spm. - Stn 73. 15 m :

2 spm. — Stn 74, 31 m : 3 spm. — Stn 7 8, 35 m :

5 spm. — Stn 85, 21 m : 1 spm. Stn 86, 29 m :

9 spm. — Stn 87, 27 m : 8 spm. — Stn 88, 34 m :

1 spm. — Stn 89, 32 m : 4 spm. - Stn 91, 30 m :

2 spm. — Stn 92, 24 m : 4 spm. — Stn 104. 24 m :

9 spm. — Stn 105, 33 m : 5 spm. Stn 107. 33 m :

7 spm. Stn 109. 16 m : 3 spm. Stn 111, 25 m :

Source : MNHN, Paris

200

WALTER O. CERNOHORSKY

4 spm. — Stn 115, 26 m : 1 spm. — Stn 119, 20 m :

3 spm. — Stn 122, 28 m : 1 spm. Stn 131, 38 m :

7 spm. — Stn 139, 45 m : 2 spm. — Stn 143, 32 m :

1 spm. Stn 147. 50-60 m : 2 spm. — Stn 149, 48 m :
1 spm. — Stn 154, 29 m : 2 spm. — Stn 155, 23 m :

3 spm. - Stn 165, 21 m : 4 spm. — Stn 167. 11m:

1 spm. — Stn 168, 10 m : 1 spm. Stn 169, 22 m :

4 spm. Stn 175. 17 m : 1 spm. — Stn 178, 20 m :

2 spm. Stn 179, 12 m : 5 spm. Stn 180, 10 m :

4 spm. Stn 187, 13 m : 3 spm. — Stn 188, 8 m :

1 spm. — Stn 192, 18 m : 2 spm. — Stn 199, 50 m :

3 spm. Stn 201, 17 m : 1 spm. — Stn 202, 13 m :

1 spm. — Stn 203, 13 m : 2 spm. — Stn 230, 35 m :

1 spm. Stn 231, 32 m : 1 spm. Stn 249, 11m:
1 spm. Stn 252, 22 m : 1 spm. — Stn 260, 23 m :

5 spm. Stn 262, 21 m : 3 spm. Stn 263, 23 m :

9 spm. — Stn 264, 19 m : 1 spm. — Stn 266, 19 m :

1 spm. — Stn 268, 24 m : 1 spm. Stn 269, 20 m :

5 spm. Stn 270, 25 m : 14 spm. — Stn 271, 22 m :
9 spm. Stn 272, 20 m : 5 spm. — Stn 274, 12 m :

1 spm. Stn 275, 19 m : 4 spm. Stn 276, 26 m :

5 spm. — Stn 277, 30 m : 4 spm. — Stn 278, 17 m :

8 spm. Stn 279, 29 m : 5 spm. Stn 285, 19 m :

1 spm. Stn 286, 28 m : 2 spm. — Stn 287, 29 m :

2 spm. — Stn 289, 23 m : 4 spm. — Stn 290, 11m:

4 spm. Stn 314. 46 m : 1 spm. — Stn 320, 70 m :

7 spm. Stn 322, 71 m : 4 spm. Stn 328, 72 m :

1 spm. Stn 331, 79 m : 3 spm. — Stn 376, 75-76 m :
1 spm. Stn 429, 95 m : 1 spm.

Distribution. — From the Red Sea and the
Persian Gulf to Japan and the Society Islands.

A ’assarius (Zeuxis) concinnus (Powys, 1835)

Nassa concima Powys, 1835 : 95.

Nassarius (Zeuxis) concinnus - Cernohorsky, 1984 :
143. pl. 28. figs 2-9 (synonymy).

Material examined. New Caledonia. Lagon :
Southwestern Lagoon : stn 143, 32 m : 2 spm.

Stn 206, 8 m : 1 spm. Stn 549, 26 m : 1 spm.

Stn 550, 24 m : 1 spm.

Eastern iMgoon : stn 672, 15-20 m : 1 spm.

Distribution. — From the Red Sea and the
Persian Gulf to Japan and the Tuamotu Archi-
pelago.

Nassarius (Zeuxis) crebricostatus (Schepman,
1911)

Nassa (Alectrvon, Acicuiina) crebricoslala Schepman,
1911 : 318.' pl. 20, figs 3a, b.

Nassarius olomea Kay, 1979 : 274, fig. 95E (new
synonym).

Nassarius (Zeuxis) crebricostatus - Cernohorsky,
1984 : 160, pl. 33, figs 3-8 (synonymy).

Material examined. - Chesterfield Islands. Musor-
stom 5 : stn 304, 22‘TO'S, 159°26'E, 385-420 m,
12 October 1986 : 1 spm. — Stn 337, 19"54'S
158°38'E, 412-430 m, 15 October 1986 : I spm.

Stn 378, 19”54' S, 158"38' E, 355 m, 20 October 1986 :
1 spm. Stn 379, 19“53'S, I58°40'E, 370-400 m,
20 October 1986 : 3 spm. Stn 381, 19°38'si
158°47' E, 620 m, 21 October 1986 : 1 spm.

New Caledonia. Biocal : stn DW 104, 21°31 ' S,
166°21' E, 375-450 m, 8 September 1985 : I spm.

Musorstom 4 : stn DW 181, 18°57'S, 163°22' E,
350 m, 18 September 1985 : I spm. Stn DW 197
18 0 5rS, 163°21'E. 550 m, 20 September 1985 :
1 spm. Stn DW 222. 22°58' S, 167°33' E, 410-440 m,
30 September 1985 : 1 spm. — Stn DW 226, 22 a 47' S,
167°22' E, 390 m, 30 September 1985 : 4 spm. —
Stn DW 227, 22"46' S, 167"20' E, 300 m. 30 September
1985 : 4 spm. Stn CC 246, 22"08' S, 167"11' E, 410-
420 m, 3 October 1985 : 6 spm. — Stn CC 247
22°09'S, 167° 13' E, 435-460 m, 4 October 1985 !
1 spm.

Distribution. - From the Red Sea to the
Tuamotu Archipelago and the Hawaiian Islands.

Remarks. This is a new record of the
species for New Caledonia.

The known bathymétrie range is extended
from 472 m to 620 m.

Some of the specimens collected were uni-
formly white or fawn in colour, while others had
the axial ribs partially or fully lined with reddish-
brown. The more slender form of N. (Z.)
crebricostatus with brown-lined axial ribs de-
scribed by Kay (1979) as N. olomea, was found
among populations of N. (Z.) crebricostatus in
New Caledonian waters, and I hâve no hésita¬
tion in placing Nassarius olomea Kay, 1979, in
synonymy of N. (Z.) crebricostatus (Schepman).

Nassarius (Zeuxis) crematus (Hinds, 1844)

Nassa cremata Hinds, 1844 : 35. pl. 9, figs 8-9.
Nassarius (Zeuxis) crematus - Cernohorsky : 129,
pl. 24, figs 1-6 (synonymy).

Material examined. New Caledonia. Lagon :
Northern Lagoon : stn 527, 59 m : 1 spm.
Southwestern Lagoon : stn 139, 45 m : 1 spm. —
Stn 147, 50-60 m : 1 spm. Stn 234, 56 m : I spm.

Stn 237, 62 m : 2 spm. - Stn 238, 50 m : 2 spm

Stn 318, 71 m : 3 spm. — Stn 321, 70 m : 1 spm. -

Stn 323, 80 m : 3 spm. Stn 325, 75 m : 3 spm. —

Stn 562, 48 m : 1 spm.

Eastern Lagoon : stn 605, 65-70 m : 2 spm. — Stn 609,
52-60 m : 1 spm. — Stn 610, 49 m : I spm. Stn 61 f.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NASSARIIDAE

201

56-57 m : 1 spm. Stn 622. 67 m : 1 spm. Stn 628,
55-56 m : 2 spm. Stn 630, 60-68 m : 4 spm. —
Stn 636, 34-40 m : 7 spm. — Stn 637, 60-65 m :
2 spm. Stn 638, 56-58 m : 2 spm. — Stn 643, 56-
66 m : 3 spm. Stn 646, 66-70 m : 1 spm. — Stn 647,
50-52 m : 7 spm. Stn 649. 64-65 m : 2 spm.

Stn 652, 55-62 m : 4 spm. — Stn 655. 35-40 m :
4 spm. - Stn 656, 30-40 m : 1 spm. — Stn 660. 48-
52 m : 1 spm. - Stn 665. 40-42 m : 4 spm. — Stn 666,
33-35 m : 3 spm. — Stn 667, 33-37 m : 1 spm. —
Stn 674, 48 m : 1 spm. — Stn 679, 29-30 m : 1 spm. —
Stn 683, 42-45 m : 2 spm. Stn 689, 46-48 m :
1 spm. Stn 691, 33-34 m : 1 spm. — Stn 692, 44-
48 m : 4 spm. Stn 694, 45-47 m : 1 spm. ; 54-55 m :

1 spm. — Stn 695, 54-55 m : 1 spm. — Stn 699, 50-
52 m : 4 spm. Stn 704, 46-58 m : 3 spm. Stn 705,
46-48 m : 2 spm. — Stn 706, 52-56 m : 1 spm. -
Stn 716, 30 m : 1 spm. Stn 722, 42 m ; 1 spm. —
Stn 723, 45 m : I spm. Stn 726, 50-51 m : 2 spm. —
Stn 727, 45-46 m : 3 spm. Stn 728, 43-47 m :

2 spm. - Stn 733, 35-38 m : 2 spm.

Distribution. From East Africa to India,
Japan and the Fiji Islands.

A 'assarius (Zeuxis) haldemanni (Dunker. 1847)
Fig. 12

Buccinum haldemanni Dunker. 1847 : 62.

Nassarius (Zeuxis) haldemanni - Cernohorsky, 1984 :
145, pl. 28, figs 10-13 (synonymy).

Material examined. New Caledonia. Lagon :
Eastern Lagoun : stn 620 : 50-52 m : 1 spm.

Distribution. - From the Red Sea to the
Philippines and the Marquesas Islands.

Remarks. — Since the publication of the mo-
nograph of Nassariidae (Cernohorsky, 1984),
specimens of N. (Z.) haldemanni hâve been
examined from Faaone, Tahiti, and lots usnm
798419, 799215 and 790397 from the Marquesas
Islands (National Muséum of Natural History,
Washington). The record of a single specimen of
this rare species from New Caledonian waters is
a new locality record, and the known bathymé¬
trie range is also extended from 40 m to 52 m.

Nassarius (Zeuxis) idyllius
(Melvill & Standen, 1901)

Nassa (Alectryon) idyllia Melvill & Standen, 1901 :
410, pl. 23,' fig. 12.

Nassarius (Zeuxis) idyllius - Cernohorsky, 1984 :
156, pl. 32, figs 8-12 (synonymy).

Material examined. New Caledonia. Lagon :
Southwestern Lagoon : stn 429, 95 m : 1 spm.

Distribution. — From the Persian Gulf to
the Philippines and the Fiji Islands.

Remarks. This is a new locality record for
the species.

Nassarius (Zeuxis) macrocephalus
(Schepman, 1911)

Nassa (Alectryon) macrocephala Schepman. 1911 :
317, pl. 20, figs 2a. b.

Nassarius (Zeuxis) macrocephalus - Cernohorsky,
1984 : 159, pl. 33, figs 1-2.

Material examined. New Caledonia. Musor-
stom 4 : stn CC 175, 18°59'S, 163"17'E, 355 m,
17 September 1985 : 20 spm.

Distribution. — From Indonesia to the
China Sea and North Australia : now New
Caledonia.

Remarks. This record is a southeastward
range extension for the species.

Nassarius (Zeuxis) siquijorensis
(A. Adams, 1852)

Nassa siquijorensis A. Adams, 1852 : 97.

Nassarius (Zeuxis) siquijorensis - Cernohorsky, 1984 :
134, pl. 25, figs 12-14; pl. 26, figs 1-5 (synonymy).

Material examined. Chesterfield Islands. Chal-
cal 1 : stn DC 33. 19 ü 45' S, 158“26' E. 205 m. 19 July
1984 : 4 spm. Stn CP 10, 20°00' S, 158°47' E. 225 m.
22 July 1981 ; 1 spm. Stn DC 63. 22"U'S,
159° 15' E. 305 m, 27 July 1984 : 7 spm. Stn DC 64.
22°11' S, 159" 15' E, 305 m, 27 July 1984 : 6 spm.

Stn DC 66, 22"26' S. 159"20' E. 320 m. 28 July 1984 :
3 spm. Stn DC 67, 22°35’ S. 159°09' E, 277 m.
28 July 1984 : 1 spm.

Musorstom 5 : stn 255, 25“15'S, 159"55'E, 280-
295 m, 7 October 1986 : 3 spm. Stn 258. 25“33' S.
159°46' E. 300 m. 8 October 1986 : 3 spm. Stn 263,
25"21'S, 159-46'E. 150-225 m, 8 October 1986 :

1 spm. - Stn 265. 25“21'S. 159"45'E. 190-260 m.
8 October 1986 : 7 spm. — Stn 266. 25°20' S,
159"46' E, 240 m, 8 October 1986 : 12 spm. — Stn 270,
24-49' S, 159"34' E, 223 m, 9 October 1986 : 5 spm. —
Stn 276, 24"49'S. 159-41'E. 258-269 m. 9 October
1986 : 3 spm. Stn 282. 24" 12'S. 159 U 32'E. 226-
230 m, 10 October 1986 : 6 spm. Stn 284. 24" 10' S.
159°33'E, 225-230 m. 10 October 1986 : 1 spm.

Stn 289. 24-02' S, 159"38' E, 273 m. 10 October 1986 :

2 spm. Stn 291, 23°08' S, 159”28'E, 300 m.

Source : MNHN, Paris

202

WALTER O. CERNOHORSKY

Il October 1986 : 3 spm. — Stn 293, 23“09' S,
159°31 ' E. 280 m. 11 October 1986 : 1 spm. Stn 294,
23°11'S. 159"30' E, 272 m, 11 October 1986 : 8 spm.
Stn 295, 23° 13' S, 159“32' E, 279 m : 11 October 1986 :
6 spm. — Stn 296. 23"13'S, I59°36'E, 278 m,
11 October 1986 : 3 spm. - Stn 312, 22“ 17'S,
159°25'E, 315-320 m, 12 October 1986 : 3 spm.

Stn 315, 22°25'S. 159°27'E, 330-335 m. 13 October
1986 : 1 spm. Stn 318, 22"27' S. I59“2r E. 330 m.
13 October 1986 : 2 spm. - Stn 320, 22°25' S,
159“ 13' E, 315 m, 13 October 1986 : 1 spm. - Stn 347,
19“39'S, 158*28' E. 260 m. 17 October 1986 : 1 spm.

New Caledonia. Musorstom 4 : stn DW 151,
19“07'S, 163“22'E, 200 m. 14 September 1985 :
I spm. Stn CC 175, I8°59'S, 163“ 17' E, 355 m,
17 September 1985 : 1 spm. Stn DW 186, 19“07' S,
163“30'E, 190 m, 19 September 1985 : 1 spm. —
Stn DW 204. 22°37' S, 167“06' E, 120 m, 27 September
1985 : 1 spm.

Lagon :

Northern Lagoon : stn 502, 190 m : I spm.
Southwestern Lagoon : stn 367, 105 m : 1 spm.

Distribution. From the Red Sea to Japan
and New Caledonia.

A as sari us ( Profundinassa) hahylonicus
(Watson, 1882)

Nassa babylonica Watson, 1882 : 366.

Nassarius ( Profundinassa) babylonicus - Cernohorsky,
1984 : 173, pl. 36, figs 1-4 (synonymy).

Material examined. New Caledonia. Biocal :
stn CP 26, 22“40' S, I66“27' E, 1 680-1 740 m, 28 August
1985 : 1 spm. - Stn CP 75, 22°19' S, 167“23' E, 825-
860 m, 4 September 1985 : 1 spm. — Stn DW 80,
20“32' S, 166“48' E. 900-980 m, 5 September 1985 :
1 spm.

Distribution. From the Gulf of Oman to
Japan and East Australia ; now New Caledonia.

Remarks. — The record of this species repre-
sents an eastward range extension from East
Australia.

Nassarius (Hitnu) pauperus (Gould, 1850)

Nassa paupera Gould, 1850 : 155.

Nassarius ( Hima) pauperus - Cernohorsky, 1984 :
176, pl. 37, figs 1-14 (synonymy).

Material examined. Chesterfield Islands. Ciial-
cal 1 : stn DC 17, 19“12' S, 158"56'e, 44 m, 17 July
1984 : 1 spm. - Stn DC 18, 19“08' S, 158“48' E, 60 m.

17 July 1984 : 1 spm. Stn DC 57, 21 "29’S
159“ 16' E. 62 m. 25 July 1984 : 1 spm.

New Caledonia. Lagon :

Huon Atoll : stn 441, 37 m : 1 spm.

Southwestern Lagoon : stn 10. 15 m : I spm. Stn 64,
15 m : 2 spm. — Stn 68, 22-40 m : 1 spm. — Stn 77,
22 m : I spm. Stn 78, 35 m : I spm. Stn 95*
14 m : 1 spm. Stn 146, 40-52 m : 1 spm. — Stn 161,
20 m : 1 spm. Stn 234, 56 m : I spm. Stn 240,

42 m : 1 spm. Stn 287, 29 m : 1 spm. Stn 290 ?,

11 m : 1 spm. Stn 348. 45 m : 2 spm. Stn 357*

77 m : I spm. Stn 384 bis, 72 m : I spm.
Eastern Lagoon : stn 603, 78-80 m : 1 spm. - Stn 619,
27-42 m : 3 spm. Stn 620, 50-52 m : 1 spm. —
Stn 621, 55-56 m : 1 spm. Stn 623, 32-40 m :
I spm. — Stn 625. 34-40 m : 1 spm. Stn 677, 32 m :
1 spm. — Stn 696, 41-57 m : 1 spm. Stn 729, 42-
45 m : 1 spm.

Distribution. From the Red Sea to South
Africa, Japan, Cook and Hawaiian Islands.

Genus CYLLENE Gray in Griffith
& Pidgeon, 1834

Cyllene concinna A. Adams, 1851
Fig. 17

Cyllene concinna A. Adams, 1851 : 205.

Cyllene concinna - Cernohorsky, 1984 : 225, pl. 51,
figs 3-5 (synonymy) ; 1988 : 84.

Material examined. Chesterfield Islands. Musor¬
stom 5 : stn 264, 25"20' S, I59°44' E, 56 m. 8 October
1986 : 4 spm.

New Caledonia. Lagon :

Southwestern Lagoon : stn 218, 15 m : 5 spm.

Distribution. From Réunion Island to
Japan ; now New Caledonia.

Remarks. Cernohorsky (1988) recorded
the species from various stations at Réunion
Island at depths ranging from 40-340 m. The
New Caledonian record is a considérable sou-
theastern range extension. C. concinna is similar
to C. pulchella Adams & Reeve, 1850, but diflers
in having a paucispiral protoconch of 1.5-2
whorls.

Cyllene fuscata A. Adams, 1851
Fig. 18

Cyllene fuscata A. Adams, 1851 : 205.

Cyllene fuscata - Cernohorsky, 1984 : 220, pl. 47
fig. 10; pl. 48, figs 1-5 (synonymy).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : NASSARI1DAE

203

Material examine!). New Caledonia. Musor-
STOM 4 : stn DW 150. 19°07' S, 163°22' E, 110 m.
14 Septembcr 1985 : 1 spm.

Distribution. From the Gulf of Oman to
East Africa and Indo-Malaysia ; now New Cale¬
donia.

Remarks. The single specimen recorded
from New Caledonian waters belongs to the
form plumbea Sowerby, 1859. This a considé¬
rable eastward range extension for the species.

Cyllene pulchella A. Adams & Reeve, 1850

Cyllene pulchella A. Adams & Reeve, 1850 : 33. pl. 10.
fig. Il-

Cyllene pulchella - Cernohorsky, 1984 : 272, pl. 49.
figs 1-8 (synonymy).

Material examined. — New Caledonia. Musor-
stom 4 : stn DW 149. 19°08' S. 163"23' E. 155 m.
14 September 1985 : 1 spm.

Distribution. — From the Red Sea to Japan
and Vanuatu.

ACKNOWLEDGEMENTS

I am grateful to Dr P. Bouchet. Muséum
national d’Histoire naturelle, Paris, for the
opportunity to examine and report upon the
family Nassariidae from New Caledonian waters.

I also would like to thank Mr P. Lozouet and
Ms V. Héros from the same institution, for their
assistance with photography and editing.

REFERENCES

Adams, A., 1851. — A monograph of Cyllene, a genus
of gasteropodous Mollusca. Proc. Zool. Soc. LonJ..
18 : 204-206.

Adams, A., 1852-1853. - Catalogue of the species of
Nassa, a genus of gasteropodous Mollusca belonging
to the family Buccinidae, in the collection of Hugh
Cuming Esq., with the description of some new
species. Proc. Zool. Soc. Lond.. 19 : 94-114 (pp. 94-
112 published 7 December 1852 ; pp. 113-114
published 29 April 1853).

Adams, A., & Reeve, L„ 1850. The Zoology of the
voyage of H.M.S. “ Samarang ”. Mollusca. Lon¬
don, 2 : 25-44, pis 10-17.

Baird, W„ in Brenchley, J. L.. 1873. — Cruise of
H.M.S. “Curaçao" among the South Sea Islands.
Section “ Shells ”, London : 432-454, pis 36-42.

Barnard, K. H.. 1959. Contributions to the
knowledge of South African Marine Mollusca.
Part 2. Gastropoda : Prosobranchiata : Rachiglossa.
Ann. S. Afr. Mus.. 45 : 1-256.

Cernohorsky, W. O., 1978. Tropical Pacific
marine shells. Sydney : 352 pp., 68 pis.

Cernohorsky. W. O., 1980. Taxonomie notes on
Polynesian Mollusca with descriptions of new spe¬
cies of Nassariidae. Rec. Auckl. Inst. Mus.. 17 : 113-
125.

Cernohorsky, W. O.. 1984. — Systematics of the
family Nassariidae (Mollusca : Gastropoda). Bull.
Auckl. Inst. Mus.. 14 : 1-356.

Cernohorsky, W. O., 1988. The Mitridae. Costel-
lariidae and Nassariidae (Mollusca : Gastropoda)
recently dredged at Reunion Island, Indian Océan,
with descriptions of new species. Rec. Auckl. Inst.
Mus.. 25 : 75-85.

Deshayes, G. P., in Belanger, C., 1832. Voyage
aux Indes Orientales par le Nord de l'Europe...
pendant les années 1825-1829... Zoologie. Mol¬
lusques. Paris : 403-440, pis 1-3.

Dunker, G., 1846. — Diagnoses Buccinorum quorun-
dam novorum. Z. Malakozool., 3 : 170-172.

— 1847. — Diagnoses Buccinorum quorundam no¬
vorum. Z. Malakozool.. 4 : 59-64.

Gould, A. A.. 1850. — Shells collected by the
United States Exploring Expédition. Proc. Bost.
Soc. Nat. Hist.. 3 : 151-156; 169-172.

Hidalgo, J. G., 1904. — Catalogo de los moluscos
testaceos de las islas Filipinas, Jolo y Marianas.
Rev. R. Acad. Cienc. Madrid, I (3) : 153-210.

Hinds. R. B.. 1844-1845. The zoology of the
voyage of H.M.S. " Sulphur” ; Mollusca. London,
1 : 1-24, pis 1-7; 2 : 25-48, pis 8-14 (1844); 3 :
49-72, pis 15-21 (1845).

Kay, E. A., 1979. — Hawaiian marine shells. Reef
and shore fauna of Hawaii. Section 4. Mollusca.
Spec. Publ. Bernice P. Bishop Mus.. 64 (4) : 1-653.

Kiener, L. C.. 1834-41. Species general et iconogra¬
phie des coquilles vivantes. Famille des Purpuri-
fères. Genre Buccin. Paris, 9 : 1-108, pis 1-31 (p. 1-
104 published in 1834; p. 105-112 in 1841).

Source : MNHN, Paris

204

WAI.TF.R O. CERNOHORSKY

Ladd, H. S., 1976. — New pleistocene Neogastropoda
from the New Hébrides. Nautilus. 90 (4) : 127-138.

Linnaeus, C., 1758. — Systema naturae per régna
tria naturae. Editio décima, reformata. Stockholm :
824 pp.

Marrat, F. P„ 1877. — On some proposed new
forms of the genus Nassa. 15 pp., 1 pl.

Melvill, J. C., & Standen. R., 1895-1896. Notes
on a collection of shells from Lifu and Uvea,
Loyalty Islands. formed by the Rev. James and
Mrs. Hadfield, with list of species. J. Conch., 8 : 84-
132, pis 2, 3 (1895); 8 : 273-315, pis 9-11 (1896).

Melvill. J. C.. & Standen, R., 1899. Report on
the marine Mollusca obtained during the first
expédition of Prof. A. C. Haddon to the Torres
Straits, in 1888-1889. J. Linn. Soc. Lond.. (Zool.).
27 : 150-206, pis 10. 11.

Melvill, J. C., & Standen, R., 1901. — The
Mollusca of the Persian Gulf, Gulf of Oman, and
Arabian Sea, as evidenced mainly through the
collections of Mr. F. W. Townsend, 1893-1900;
with descriptions of new species. Proc. Zool. Soc.
Lond.. 1901 (2) : 327-460. pis 21-24.

Melvill, J. C.. & Standen, R.. 1903. Descriptions
of sixty-eight new Gastropoda from the Persian
Gulf, Gulf of Oman, and North Arabian Sea,
dredged by Mr. F. W. Townsend, of the indo-
European Telegraph service, 1901-1903. Ann. Mag.
Nat. Hist.. (7) 12 : 289-324, pis 20-23.

Powys, W. L., 1835. — Characters of new species of
shells collected by Mr. Cuming. Proc. Zool. Soc.
Lond.. 3 : 93-96.

Quoy, J. R. C.. & Gaimard, J. P.. 1833. Voyage
de découvertes de l'Astrolabe exécuté par ordre du
roi, pendant les années 1826-1829. sous le comman¬
dement de M. J. Dumont d'Urville. Zoologie, 2.
Paris ; 321-674, pis 24-45 bis.

Reeve, L., 1853-1854. — Conchologia Iconica, 8;
monograph of the genus Nassa. London : pis 1-29,
text.

Rjcher de Forges, B., 1990. Les campagnes
d’exploration de la faune bathyale dans la zone
économique de la Nouvelle-Calédonie. In : A. Cros-
nier (ed.). Résultats des campagnes Musorstom,
Volume 6. Mém. Mus. nain. Hist. nat., (A) 145 : 9-
54.

Richer de Forges, B., & Barcibant, G., 1985. Le
lagon Nord de la Nouvelle-Calédonie et les atolls de
Huon et Surprise. Orstom, Centre de Nouméa.
Rapports scientifiques et techniques, 37 : 1-23.

Richer de Forges, B., et al.. 1987. Le lagon sud-
ouest de la Nouvelle-Calédonie. Observations préa¬
lables à la cartographie bionomique des fonds
meubles. Orstom, Centre de Nouméa. Rapports
scientifiques et techniques. Sciences de la Mer, 45
1 - 110 .

Schepman, M. M.. 1911. The Prosobranchia of the
Siboga Expédition. Part 4. Rachiglossa. Siboga-
Expedition, 49d : 247-363, pis 18-24.

Watson, R. B., 1882. Mollusca of H.M.S. “Chal¬
lenger" Expédition. Part 13. J. Linn. Soc. Lond.,
(Zool.) 16 : 358-372.

Source : MNHN, Paris

JLTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSUL1

8

Mollusca Gastropoda : Eumitra récentes de la région
néo-calédonienne et Charitodoron fossiles
de l’Oligocène supérieur d’Aquitaine (Mitridae)

Pierre LOZOUET

Muséum national d'Histoire naturelle
Laboratoire de Biologie des Invertébrés marins et Malacologie
55, rue Buffbn
75005 Paris

RÉSUMÉ

Des espèces récentes A'Eumitra sont décrites pour la
première fois ; elles proviennent du bathyal de la région néo-
calédonienne : E. caledonica sp. nov., E. apheles sp. nov., E.
imbricata sp. nov. et E. richeri sp. nov. Les Eumitra n'étaient
connues que du Miocène inférieur de Nouvelle-Zélande et du
Miocène/Pliocène d'Australie. La dispersion peut expliquer
la distribution du genre. Deux espèces fossiles provenant de

paléocommunautés bathyales de l'Oligocène supérieur d'Aqui¬
taine sont pour la première fois rapportées au genre Charito¬
doron. Les trois espèces actuelles de Charitodoron sont
limitées au bathyal supérieur d'Afrique du Sud. Charitodoron
et Eumitra morphologiquement proches n'apparaissent pas
directement reliés.

ABSTRACT

Mollusca Gastropoda : Rccent Eumitra from New Calcdo-
nian région and fossil Charitodoron from upper Oligocène of
Aquitaine (Mitridae).

The first Recenl species of Eumitra are described from
deep-water in the New Calcdonian région : E. caledonica
sp. nov. (Southern New Calcdonia). E. apheles sp. nov.
(Northern New Caledonia), E. imbricata sp. nov. (Coral Sca.
Lansdowne-Fairway) and E. richeri sp. nov. (Coral Sea.
Mcllish Reef). A SEM photograph of the radula is included.
Fossil Eumitra arc restricted to lower Miocène of New
Zealand and Miocene/Pliocene of Australia. Dispersai is
advocated to explain Eumitra distribution. For the first time

fossil species from Upper Oligocène of Aquitaine Basin
(Southwestcrn France) are referred to Charitodoron, an
atypical member of the Mitridae : C. tauzini sp. nov. and C.
cancellants (Saubade, 1969). The three Recenl Charitodoron
are confined to the bathyal zone of South Africa, fossil
Oligocène species hâve been collected from a bathyal palaeo-
community. In spitc of columellar similarities. peculiar
development of columellar folds ( Eumitra ) or edcntulous
columella (Charitodoron). these two gênera arc probably not
closely related. In a paleobiogeographic discussion two key
events are citcd to explain the beginning of many marine
disjunctions : Upper Eocene/Lowcr Oligocène crisis and
closing of Tethys in Upper Oligoccne/Lower Miocene.

Lozouht. P., 1991. Mollusca Gastropoda : Eumitra récentes de la région néo-calcdonienne et Charitodoron fossiles de l'Oligocène
supérieur d'Aquitaine (Mitridae). In : A. Crosnier & P. Bouchet (eds). Résultats des Campagnes Musorstom. Volume 7. Mêm. Mus. nain.
Hisl. nul.. (A). 150 : 205-222. Paris ISBN : 2-85653-180-6.

Public le 20 mars 1991.

Source : MNHN, Paris

206

PIERRE LOZOUET

INTRODUCTION

L'une des caractéristiques des Milridae est la
présence de plis columellaires saillants. Deux
exceptions sont cependant à signaler : le genre
Charitodoron (limité aux côtes de l’Afrique du
Sud) dont les espèces ne présentent aucune
plication columellaire et le genre Eumilra (uni¬
quement connu du Néogène australo-zélandais)
chez lequel les plis sont faibles. Se basant sur ce
caractère. Cernohorsky (1970) a proposé une
filiation Eumitra-Charitodoron. Les Charitodoron
se seraient séparés des Eumilra au cours du
Pliocène. Or, deux espèces présumées de Charito¬
doron ont été récoltées dans l’Oligocène supé¬
rieur d’Aquitaine. D’autre part quatre espèces
d 'Eumitra ont été draguées lors des campagnes
Biocal (Chef de Mission : C. LÉvi), Musorstom
4 (Chef de Mission : B. Richer de Forges) et
Corail 2 (Chef de Mission : B. Richer de

Forges) dans le bathyal supérieur de Nouvelle-
Calédonie et de la mer du Corail (banc Lans-
downe-Fairway, Mellish Reef). Ces découvertes
obligent à repenser le schéma de filiation de
Cernohorsky et suscitent des interrogations
biogéographiques.

Dans le cours du texte, différentes abréviations
ont été utilisées :

ipm : Institut de Paléontologie du Muséum
national d’Histoire naturelle, Paris
mnhn: Muséum national d’Histoire naturelle,
Paris

nm : Natal Muséum, Pietermaritzburg
smf : Natur-Museum Senckenberg, Francfort
ams : The Australian Muséum, Sydney.

ÉTUDE SYSTÉMATIQUE
Famille des MITRIDAE
Genre CHARITODORON Tomlin, 1932

Espèce-type. Columbella harhara Thiele,
1925.

Les premières espèces décrites furent classées
dans les Bucçinidae ( Columbella agulhasensis
Thiele, 1925 : 173, pl. 18 fig. 20; Columbella
barbara Thiele, 1925 : 173, pl. 18 fig. 22). Tomlin
(1932) redécrit ces espèces (dans l’ignorance du
travail de Thiele, 1925), en ajoute une troisième
(Charitodoron thulia ) et crée le genre Charitodo¬
ron qu’il classe dans les Buccinidae. Tomlin
(1943) décrit une quatrième espèce ( Charitodoron
pasithea) et Barnard (1959) une cinquième qu’il
place dans les Mitridae [Mitra (Dibaphus) bathy-
bius] à la suite de l'examen de la radula.
Barnard (1960) examine la radula de Charitodo¬
ron thalia qui s'avère identique à celle de l’espèce

bathybius. 11 conclut au classement de Charitodo¬
ron dans les Mitridae et propose des rectifica¬
tions de nomenclature. Enfin, Cernohorsky
(1970, 1976) réexamine les différentes espèces de
Charitodoron.

Le genre Charitodoron ne comprend finale¬
ment que trois espèces actuelles récoltées par
dragages entre 150 m et 1 300 m, le long des
côtes de l’Afrique du Sud (de Cape Point à East
London) :

Charitodoron barbara (Thiele, 1925) ( = euphro-
syne Tomlin, 1932)

Charitodoron agulhasensis (Thiele, 1925) (= aglaia
Tomlin, 1932)

Charitodoron thalia Tomlin, 1932 (= pasithea
Tomlin, 1943; bathybius Barnard, 1959).
Outre sa columelle sans pli, Charitodoron se

Source : MNHN, Paris

MOLLUSCA GASTROPODA : EU MIT RA ET CHARITODORON

207

Fig. 1-4. — Charilodorun de l'Oligocène supérieur. 1-3.
Charitodoron tauzini (1. holotypc. Si-Étienne-d'Orthe. ii*m
r 3074. -- 2, Peyrehorade, MNHN-Malacologie. 3. para-
type, St-Étienne-d'Orthe. ipm R53075). 4. Charitodoron

cancellatus, MNHN-Malacologie. St-Étienne-d'Orthe (Oligo¬
cène supérieur).

caractérise par une protoconque assez grosse et
courte (indiquant un développement non-planc-
totrophe) et un labre fin non épaissi.

Charitodoron tauzini sp. nov.

Fig. 1-3, 7, 18, 33-34, 38

Localité type. St-Étienne-d’Orthe, bassin
d’Aquitaine (France).

Étage type. — Oligocène supérieur, marnes à
Miogypsinoides.

Matériel type. — Holotypc : ipm r53074
(coll. Tournouër). Paratypes : St-Étienne-d'Orthe
(ipm r53075 à 53082, 8 ex., coll. Tournouër ;
Muséum de Bordeaux, 2 ex., coll. Degrange-
Touzin) ; St-Étienne-d'Orthe : ruisseau de l’Église
(10 ex.), Troun (4 ex.) (MNHN-Malacologie, coll.
Lozouet).

Autre matériel. — Peyrehorade : Peyrère
(2 ex.. MNHN-Malacologie, coll. Lozouet).

Description. - Coquille de petite taille, fusi¬
forme, élancée, composée de six tours trois-
quarts de téléoconque convexe à suture marquée.
L’apex est mammillé peu saillant ; il se termine
par de fines stries longitudinales auxquelles suc¬
cèdent assez insensiblement les premières cos-
tules axiales. Entre ces dernières se développent
rapidement des cordons spiraux. L’ornementa¬
tion est ainsi constituée sur les trois premiers
tours post-larvaires, de côtes axiales au nombre
de 19-20 et de cordons spiraux (6-7), plus larges
que leur intervalle, dont l’intersection forme de
petits tubercules. Cette ornementation des pre¬
miers tours de téléoconque tend par la suite à
devenir obsolète. Sur les derniers tours de la
coquille ne subsistent que deux ou trois sillons
spiraux bordant la suture et des fortes stries
d'accroissement. Sur le dernier tour qui occupe
un peu moins des 2/3 de la hauteur totale se
développent en outre, sur sa base, de forts filets
spiraux décussés par les stries d’accroissement.
L'ouverture allongée est plus grande que la spire,
le canal siphonal est légèrement recourbé et
possède une fasciole basale ; la columelle est lisse
avec un fin inductura. Le labre fin, lisse intérieu¬
rement, se termine par une échancrure siphonale

Source : MNHN, Paris

208

PIERRE LOZOUET

parfaitement distincte et son canal anal est bien
marqué.

Protoconque : globuleuse, d'un tour et demi
convexe, lisse. Le passage à la télèoconque n'est
pas nettement matérialisé.

Dimensions (holotype) : hauteur 18,3 mm ;
diamètre 5,8 mm.

Discussion générique et spécifique. — Chari-
todoron tauzini sp. nov. se distingue de Parvi-
sipho cancellatus Saubade, 1969 (Fig. 4, 19, 31-
32) des mêmes dépôts, par sa faible sculpture,
obsolète sur les derniers tours, et sa protoconque
moins acuminée.

L'examen des figures de C. tauzini (Fig. 1-3)
montre deux formes de galbe différent. La forme
de Peyrère est plus allongée et les derniers tours
de la coquille ne présentent pas de sillons en
bordure de la suture. Une variabilité comparable
existe chez les Charitodoron actuels notamment
avec les espèces Charitodoron agulhasensis et
C. agi nia que Cernohorsky (1976) considère
comme des synonymes.

L'espèce tauzini sp. nov. est morphologique¬
ment très comparable à Charitodoron harbara et
C. agulhasensis des côtes d’Afrique du Sud.
L'espèce oligocène est toutefois de taille moindre
et son canal anal est net. D’autre part, le canal
siphonal est plus recourbé chez tauzini , peut-être
plus proche de celui des espèces du genre Metula
(Buccinidae) que des Mitridae (généralement
court et subdroit). Il est à souligner que Parvi-
sipho cancellatus qui semble indissociable généri¬
quement de C. tauzini, présente une sculpture
proche de celle des Metula. Toutefois, les espèces
de Metula H. & A. Adams, 1853 (espèce-type :
Buccinum clathratum Adams & Reeve, 1850)
semblent avoir, toutes, un péristome épaissi
pouvant former une varice et souvent le bord
interne est finement denticulé. Je note aussi que,
chez Metula, le canal siphonal (Fig. 10) ne
présente pas de fasciole. or les espèces tauzini et
cancellatus en possèdent une très nette. Charito¬
doron barbara et agulhasensis ont aussi un canal
muni d’une fasciole (Fig. 9), celle-ci est cepen¬
dant moins prononcée que chez C. tauzini.

Le genre Parvisipho Cossmann, 1889 (espèce-
type : Fusus terebralis Lamarck. 1804; Éocène
moyen, Lutétien du bassin de Paris), dans lequel

Saubade (1969) a rangé l'espèce cancellatus, est
aussi un Buccinidae (au sens de Ponder et
Waren, 1988). Chez l'espèce-type (Fig. 41-42), le
labre est bordé extérieurement, finement denti¬
culé intérieurement et le canal siphonal court est
dépourvu de fasciole (le concept de Parvisipho de
Le Renard, 1989, n'est pas adopté car ce dernier
en exclut l'espèce terebralis).

Janssen (1979) a rangé aussi dans le genre
Parvisipho une espèce assez proche de C. tauzini
et cancellatus (Fusus scrobiculatus Boll, 1851 ;
Oligocène supérieur d'Allemagne). Fusus scrobi¬
culatus possède une spire plus élevée, séparée par
des tours canaliculés et une sculpture à base de
forts cordons spiraux, très différente de celle de
C. tauzini. Son canal siphonal, plus étroit,
nettement individualisé du reste du dernier
tour (d'après l'unique individu observé, smf
250349/1), se distingue nettement de celui de
C. tauzini (voir Fig. 6). Il ressemble un peu à
celui du Turridae Pusionellu (genre endémique à
la province ouest-africaine). Ce dernier genre
compte des espèces morphologiquement assez
proches de C. tauzini, mais avec un canal
siphonal plus étroit et allongé.

Dans cette discussion, trois familles de Neo-
gastropoda ont été évoquées (Buccinidae, Mi¬
tridae et Turridae) avec respectivement les genres
Metula/Parvisipho, Charitodoron et Pusionella.
Le classement des espèces tauzini et cancellatus
dans le genre Charitodoron, Mitridae atypique,
paraît le mieux convenir. La principale diver¬
gence réside dans la forme du canal siphonal.
Des différences toutefois aussi nettes ont été
notées avec le canal de Metula, Parvisipho et
Pusionella.

Remarques. Dans les collections du Mu¬
séum de Bordeaux (coll. Di-grange-Touzin)
mais aussi dans celle de la Faculté de Bordeaux-
Talence (coll. Reyt), C. tauzini est identifié sous
le nom de Linderia aturensis Peyrot. À ma
connaissance le genre Linderia et l’espèce aturen¬
sis sont des noms manuscrits.

Étymologie. Dédiée à l’Abbé Tauzin,
premier récolteur à St-Étienne-d'Orthe probable¬
ment autour de 1860-1870.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : EU MITRA ET CHARITODORON

209

Fig. 5-10. Canal siphonal : 5, Pusionella aculeiformis (Lamarck, 1822), Côte d'ivoire. Région d'Abidjan, dragages (mnhn.
coll. Marche-Marchad). 6. Fusus scrobiculatus. Oligocène supérieur (smf 250349/1, coll. Gorges). 7. Charitodoron
tauzini, holotype. Oligocène supérieur (ipm r53074, coll. Toumouër). 8, Pusionella milleti (Pelit. 1851), Gambie,
S. Cape Bald (mnhn). 9. Charitodoron aguthasensis, Afrique du Sud, Cape St Blai/e, estomac de Congiopodes
(nm a4054). 10, Metula africana Bouchet, 1988, Sénégal, St Louis, 300-600 m, (mnhn, leg. M. Pin). Échelle = 10 mm.

Source : MNHN, Paris

210

PIERRE LOZOUET

Genre EU MIT R A Tate, 1889

Espèce-type. Mitra alokiza Tenison-Woods,
1880. Miocène moyen du sud-est de l'Australie.

Eumitra se caractérise essentiellement (Cerno-
horsky, 1970) par la faiblesse de ses plis colu-
mellaires. Un seul pli est généralement bien
développé. C’est sur la base de ce caractère que
Finlay (1926) a créé le genre Diplomitra pour
des espèces du Miocène inférieur de Nouvelle-
Zélande. Tel que le comprend Cernohorsky, le
genre Eumitra englobe Diplomitra. Cependant,
Mitra alokiza sensu Tate (1889) possède 5 plis
columellaires décroissants. Cernohorsky (1970)
précise que ceux qu'il a observé ont un pli
proéminent et l'indice d'un second, mais il admet
la présence d'individus pouvant en posséder
davantage.

La coquille des Eumitra présente une faible
sculpture spirale, toutefois les premiers tours
peuvent présenter des côtes axiales notamment
chez Eumitra dictua (Tenison-Woods, 1880) du
Miocène australien.

Le genre Eumitra semblait limité au Néogène
australo-zélandais (Miocène inférieur à Pliocène
inférieur) où il ne compte qu’un petit nombre
d'espèces. Toutefois, P. Maxwell (comm. pers.)
me signale la similitude entre Eumitra alokiza et
le Mitridae du Miocène moyen de la Paratéthys
rapporté à Concilia scrobiculata (Brocchi, 1814)
(espèce décrite du Pliocène d’Italie, Rossi Ron-
chetti, 1955, fig. 131). J'ai examiné du groupe de
Concilia scrobiculata : une quarantaine d’exem¬
plaires du Miocène moyen du bassin de Vienne
(Baden), cinq exemplaires du Miocène de Sau-
brigues (France, Landes) et différents lots du
Pliocène d’Italie. Effectivement, quelques échan¬
tillons rappellent E. alokiza. Il existe une varia¬
bilité importante de la sculpture, de la force et
du nombre des plis columellaires mais ceux-ci
restent cependant compris entre 3 et 5 (Pour
Cernohorsky, 1970, Concilia présente de 3 à
6 plis columellaires). Concilia scrobiculata sensu
Hôrnes (1851 : PI. 10, fig. 14-18) ne peut donc
pas être rapportée au genre Eumitra. Cette
espèce suscite cependant, en raison de sa ressem¬
blance avec Eumitra alokiza, des interroga¬
tions quant à l'origine d'Eumitra et des rela¬
tions éventuelles Conciliai Eumitra. Notons que

le groupe des Concilia (s. s.) Swainson, 1840
(espèce-type : isabella Swainson, 1831 ; de l’Indo-
Ouest-Pacifique), tel que le comprend Cerno¬
horsky (1970), apparaît fondé uniquement sur
les caractères de la coquille. Aucune radula du
groupe ne semble avoir été examinée.

Eumitra caledonica sp. nov.

Fig. 11, 43-46

Matériel type. Nouvelle-Calédonie (sud de
l'île des Pins). Biocal : Holotype (mnhn), stn 61,
24”11,67' S-I67°31,37' E, 1 070 m, coquille vide.
Paratypes, 3 exemplaires (mnhn) : 1 ex. de la
même station que Phoiotype (stn 61); stn 70,
23°24,70'S-167°53,65'E, 965 m, 1 ex.; stn 62,
24° 19' S-167°49' E, 1395-1410 m. 1 ex. Maté¬
riel récolté par P. Bouchet, B. Métivier et
B. Richer de Forges à bord du N. O. “ Jean
Charcot ",

Description. — Coquille de taille moyenne,
fusiforme, élancée. La téléoconque se compose
de 5 tours 1/4 convexes séparés par une suture
bien marquée. Apex peu distinct de la téléo¬
conque. La sculpture se limite à de nombreuses
mais faibles stries axiales, entrecoupées de stries
spirales obsolètes. Toutefois, il existe, sur le
premier tour et demi de la téléoconque, une
sculpture de côtes axiales, légèrement renflées en
bordure de la suture et formant un petit tuber¬
cule. Ouverture ovoïde ; avec un labre fin, lisse
intérieurement, de direction légèrement opistho-
cline. Le bord pariétal et columellaire est recou¬
vert par un mince inductura. La columelle
présente en sa partie médiane deux faibles plis
transverses de même force. Le canal siphonal est
court, subdroit et assez largement ouvert.

Coloration uniformément blanchâtre.

Protoconque : d’environ 2 tours 1/2 convexes
et nucléus assez petit. Le passage à la téléo¬
conque est peu distinct.

Dimensions (holotype) : hauteur 35 mm ; dia¬
mètre 10,3 mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : EUM/TRA ET CH A R1TODORON

211

Fig. 11-19. Protoconques : 11, Eumiira calédonien, hololypc. Nouvelle-Calédonie, Biocal, stn 61. 12, Eumitra apheles

holotype, Nouvelle-Calédonie, Musorstom 4. stn 159. 13. Eumiira imbricata, holotype, mer du Corail. Lansdowne-

Fairway, Corail 2, stn 14. 14. Pusionella aculeiformis. Côte d’ivoire, région d'Abidjan. — 15, Eumiira apheles,

paratype, Nouvelle-Calédonie, Musorstom 4. stn 159. — 16, Charitodoron agulhasensis. Afrique du Sud, Cape

St Blaizc (NM A4054). — 17. Fusus scrohiculalus,
holotype. Oligocène supérieur (ipm r53074). 19.

St-Etienne-d’Orthe. Échelle = 1 mm.

Eumitra apheles sp. nov.

Fig. 12, 15, 20. 47-50

Matériel type. Nord de la Nouvelle-Calé¬
donie (Grand Passage), Musorstom 4 : Holotype
(mnhn), stn 159, 18°45,90' S-163°15,60' E, 600 m.
Paratypes, 11 exemplaires (mnhn, nmnz, ams) :

Oligocène supérieur (smf 250349/1). 18. Charitodoron lauzini.

cancellalus. Oligocène supérieur, bassin d’Aquitaine,

2 ex. de la même station que l’holotype ; stn 161,
18"38,80' S-163° 10,60' E, 565 m, 2 ex. ; stn 168,
18'’48,20' S-163° 10,80' E, 720 m, 7 ex. Matériel
récolté par P. Bouchet et B. Richer de Forges
à bord du N. O. “ Vauhan

Description. — Coquille de petite taille, fusi¬
forme, élancée, composée de 4 tours 1/2 convexes

Source : MNHN, Paris

212

PIERRE LOZOUET

à suture bien marquée. La sculpture se compose
de nombreuses mais faibles stries axiales extrê¬
mement fines, recoupées par quelques stries
spirales. Sur la base de la coquille, au niveau du
canal siphonal, on distingue 4-5 cordons spiraux
très obsolètes. Ouverture ovoïde ; labre de direc¬
tion presque orthocline, très légèrement épaissi,
lisse intérieurement. Columelle recouverte par un
inductura excessivement fin ; elle porte, en sa
partie médiane, deux faibles plis inégaux ; le
premier (adapical) plus fort semble perpendicu¬
laire à l'axe de la columelle, le second (abapical)
obsolète apparaît plus parallèle. Le canal sipho¬
nal est court, presque droit, assez largement
ouvert.

La coquille présente une coloration blan¬
châtre.

Protoconque : d’environ 2 tours convexes, à
nucléus assez petit ; peu distincte de la téléo-
conque.

Dimensions : (holotype) : hauteur 15,6 mm ;
diamètre 5,6 mm.

Radula : Rachiglosse de type Mitridae. Dent
centrale petite, avec 5-6 cuspides réparties symé¬
triquement de part et d'autre d'une cuspide
centrale. Dents latérales grandes, allongées,
munies de 9-10 cuspides beaucoup plus fortes
que celles de la dent centrale.

Discussion. — Eumitra cipheles se sépare
d'E. caledonica par sa protoconque et sa taille
plus petite. Eumitra aplieles est aussi totalement
dépourvue de côte axiale alors qu’il existe chez

Fig. 20. — Radula d'Eumitra aplieles, Musorstom 4, stn 168.
720 m. Échelle = 100 |xm.

E. caledonica une sculpture de côtes axiales
faibles, mais nette sur le premier tour et demi.

Les Eumitra fossiles décrites soit par Mar¬
shall (1918). Powell et Bartrum (1929) pour
la Nouvelle-Zélande, soit par Tenison-Woods
(1880), Tate (1889) pour l'Australie, appa¬
raissent très distinctes. Les espèces les plus
proches (£. uniplica Tate, 1889, du Miocène
moyen d’Australie ; Eumitra waitemataensis Powell
& Bartrum, 1929, du Miocène inférieur, Otaian)
possèdent une suture beaucoup plus canaliculée
et une téléoconque à tours moins convexes.

ÉTYMOLOGIE. Lisse, non raboteuse (du grec :
àçeXvjç).

Eumitra imbricata sp. nov.

Fig. 13, 21, 56-58

Matériel type. Mer du Corail, Lans-
downe-Fairway, Corail 2. Holotype unique
(mnhn). Stn 14, 21 °00,69' S-160°57,18' E, 650-
660 m.

Description. Coquille de taille moyenne,
fusiforme, élancée, composée de 4 tours 3/4.
Les deux premiers tours de téléoconque, peu
convexes, sont séparés par une suture fortement
canaliculée, leur donnant un aspect imbriqué.
Les tours suivants présentent un profil assez
régulièrement convexe et la suture est moins
marquée. Le singularisme des deux premiers
tours est encore renforcé par une sculpture
vigoureuse, composée de 14-15 côtes axiales
assez fortes, moins larges que leur intervalle.
Cette sculpture disparaît après le deuxième tour
et on ne distingue que des stries d’accroissement,
plus ou moins marquées, et des filets spiraux très
obsolètes. Sur la base de la coquille, au niveau
du canal siphonal, on observe une dizaine de
faibles cordons spiraux. L'ouverture est ovoïde.
Le labre, d’après les stries d'accroissement, devait
être de direction orthocline. La columelle est
recouverte par un faible inductura. Elle porte, en
sa partie médiane, deux forts plis inégaux. Le
premier (adapical) est plus marqué et semble plus
nettement perpendiculaire à l'axe de la columelle.
Le canal siphonal est court, presque droit, assez
largement ouvert.

La coquille présente une coloration blan¬
châtre.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : EU MITRA ET CH A R/TODORON

213

Protoconque : bien individualisée de la téléo-
conque, de 2 tours 1/2 à nucléus assez petit.

Dimensions : (holotype) : hauteur 15,6 mm ;
diamètre 5,2 mm.

Discussion. Eumitra imbricata sp. nov. se
distingue immédiatement d'E. apheles et E. calé¬
donien par la vigueur de l'ornementation des
premiers tours, sa suture fortement canaliculée,
l'aspect imbriqué des tours et la force des deux
plis columellaires.

La forme générale d'E. imbricata rappelle
Eumitra waitemataensis (Powell & Bartrum, 1929)
du Miocène inférieur de Nouvelle-Zélande dont
elle se sépare par la sculpture de ses premiers
tours.

Etymologie. Du latin imbricata, allusion à
l’étagement des tours.

FlG. 21. Eumitra imbricata. Holotype. Échelle = 5 mm.

Eumitra richeri sp. nov.

Fig. 51-55

Matériel type. Mer du Corail, SW de Mel-
lish Reef, Corail 2. Holotype (ams), stn 172, 18°
25,55' S-155° 12.82' E, 1 100 m. Paratypes (mnhn) :
2 ex. de la même station que l'holotype.

Description. — Coquille de taille moyenne,
fusiforme, élancée, composée d'un peu plus de
4 tours 3/4 de téléoconque séparés par une suture
bien marquée. La sculpture se compose d'une
dizaine de côtes axiales, restreintes aux deux
premiers tours de téléoconque, et d'assez nom¬
breux cordons spiraux limités au dernier tour. Il
existe aussi de nombreuses stries d’accroisse¬
ment. L'ouverture est ovoïde avec un labre
presque orthocline. Le bord pariétal et columel-
laire est recouvert par un faible inductura. La
columelle présente, en sa partie médiane, deux
plis columellaires inégaux, le premier (adapical)
est plus fort. Le canal siphonal est court, sub¬
droit, assez largement ouvert.

Coquille blanchâtre.

Protoconque : de 2 tours 1/4, bien individua¬
lisée de la téléoconque.

Dimensions : (holotype) : hauteur 22,5 mm ;
diamètre 7,7 mm.

Remarques. — Les deux paratypes ont des
plis columellaires plus faibles, en particulier
l'exemplaire de la figure 51.

Discussion. — La présence de cordons spi¬
raux bien marqués sépare cette espèce des autres
Eumitra calédoniennes et en particulier d'Eu-
mitra apheles. Eumitra caledonica qui présente
une sculpture proche sur le premier tour est de
taille supérieure et sa protoconque est plus
grosse.

Étymologie. Dédiée à Bertrand Rk her de
Forges.

Analyse biométrique des Eumitra

Afin de compléter les observations précé¬
dentes, 6 mesures et 3 rapports ont été effectués
puis traités par une analyse factorielle des corres¬
pondances. Ces neuf variables sont (Fig. 22) :

1 Hauteur de la coquille = hau

2 — Diamètre maximum = dma

3 Hauteur de la spire = hsh

4 — Largeur de la spire = lgs

Source : MNHN, Paris

214

PIERRE LOZOUET

5 — Hauteur de l'ouverture = hto

6 — Largeur de l’ouverture = lgo

7 Rapport hau/dma = hdm

8 — Rapport hsp/lgs = hsl

9 Rapport hto/lgo = hol

Fig. 22. — Mesures utilisées chez Eumitra.

Il convient de souligner l'hétérogénéité et la
faiblesse de l’échantillonnage (20 spécimens dont
17 ont pu être mesurés). Ainsi, la population la
moins médiocre ( Eumitra apheles) comprend des
individus très jeunes (hau = 9,4 mm) à labre
non formé et d'autres, comme l’exemplaire de la
figure 50 (hau = 16,9 mm), qui présentent diffé¬
rentes traces d’accidents de croissance.

L’axe 1 explique a lui seul 91 % de l’inertie
totale contre 2,2 % pour l’axe 2 et est relative¬
ment monopolaire. L'inertie du nuage des varia¬
bles est due aux fortes contributions des rapports
(hol, hdm). Il oppose ces dernières aux hauteurs
de la coquille et de la spire (hau, hsp). Ce qui se
traduit par une opposition entre les grands indi¬
vidus d 'Eumitra caledonica (c3, cl) et les jeunes
individus d'E. apheles (a 10, a9, a8, a3). Il
apparaît sur le graphe factoriel (Fig. 23) que les
quatre espèces ne se chevauchent pas. Cependant
si la discrimination est nette entre E. caledonica
et E. apheles , elle l’est beaucoup moins entre E.
apheles\E. imbricata d’une part, et E. caledo-
nica/E. richeri d’autre part.

Remarques sur les Eumitra néo-calédoniennes

Les caractéristiques sculpturales, bien que
ténues, ont conduit à distinguer quatre espèces.
L’absence (Eumitra apheles) ou la présence de
côtes axiales sur les premiers tours (E. caledo¬
nica. E. richeri, E. imbricata) ont été considérées

Fig. 23. — Analyse factorielle des correspondances des Eumitra. Projection des variables sur le plan factoriel 1-2. a =
E. apheles. c = E. caledonica. i = E. imbricata, R = E. richeri.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : EUMITRA ET CHAR1TODORON

215

comme significatives. Les côtes axiales sont
fortes chez Eumitra imbricata, plus faibles chez
E. caledonica et E. richeri. Chez cette dernière
espèce, le dernier tour présente une sculpture
spirale beaucoup mieux marquée. L'analyse bio¬
métrique n'infirme pas la séparation des Eumitra
de la région calédonienne en quatre espèces. Elle
conduit cependant à un rapprochement E. cale¬
donica/E. richeri d'une part, E. apheles/E. imbri¬
cata d'autre part et montre une discrimination en
fonction de la taille. La présence ou l'absence de
sculpture sur les premiers tours indique claire¬
ment que cette discrimination ne peut être
imputée à un phénomène de modification de
forme en fonction de la croissance. On ne peut
considérer, par exemple, qu Eumitra apheles
regroupe des jeunes E. caledonica.

Tout ceci suggère que nous nous trouvons en

présence de quatre unités proches, mais généti¬
quement séparées. Il convient de rappeler que :

— les protoconques des quatre Eumitra indiquent
un développement larvaire sans phase pélagique
limitant donc les possibilités d'échanges entre les
populations ;

— elles ont été draguées dans des fonds de 500 à
1 000 m en position d'isolement géographique
(Fig. 24).

Bien qu'il existe une continuité bathymétrique
entre Eumitra apheles (nord de la Nouvelle-Calé¬
donie) et Eumitra caledonica (sud de l'île
des Pins), ces deux espèces sont celles chez
lesquelles les différences, non ornementales,
apparaissent les plus tranchées. En revanche,
les espèces géographiquement les plus éloignées
l'une de l'autre (E. caledonica et E. richeri) sont
proches sur le diagramme factoriel (Fig. 23).

Source : MNHN, Paris

216

PIERRE LOZOUET

CONCLUSION ET DISCUSSION PALÉOBIOGÉOGRAPHIQUE

L'hypothèse phylétique de Cernohorsky (1970)
doit être réexaminée pour au moins deux rai¬
sons :

— Eumitra renferme au Miocène des espèces
proches morphologiquement de Mitra (s. s.). En
particulier, l'espèce-type alokiza présente parfois
5 plis columellaires. Les Charitodoron seraient,
dès l’Oligocène supérieur, dépourvus de plis.

la radula à'Eumitra apheles (Fig. 20) est bien
de type Mitridae mais se distingue nettement de
celle des autres Mitridae, y compris de celle de
Charitodoron thalia (Barnard, 1959, fig. 11;
Cernohorsky, 1976 pl. 251), par sa dent centrale
à petites cuspides.

Ces remarques suggèrent que la perte de la
plication columellaire chez Charitodoron et sa
faiblesse chez Eumitra ont été acquises indépen¬
damment et ne sont pas dues à une ascendance
directe.

Les Eumitra fossiles ont été décrites de faciès
littoraux néogènes d’après les travaux de Tate
(1889) pour l'Australie et Powell et Bartrum
(1929) pour la Nouvelle-Zélande. La localisation
au bathyal supérieur (500 à 1 000 m, Fig. 24) de
Nouvelle-Calédonie et de la mer du Corail des
quatre seules espèces actuellement connues est
donc à souligner. Elle illustre une fois encore le
conservatisme de cette tranche bathymétrique.
La localisation bathymétrique des Charitodoron
actuels (bathyal essentiellement ; moyenne des
profondeurs relevées : 751 m) est conforme à
celle des deux espèces fossiles oligocènes recon¬
nues, ainsi que le montrent les reconstitutions
paléoécologiques (Lozouet, 1986).

La famille des Mitridae apparaît vers la fin du
Crétacé et les genres Charitodoron et Eumitra ne
s’individualisent vraisemblablement pas avant le
Paléogène supérieur.

L’éclatement de la Tasmantis et la dérive de
ses constituants sont donc trop anciens pour
expliquer par un déplacement passif la répar¬
tition actuelle d’ Eumitra. La dispersion rend
mieux compte de ce phénomène. Eumitra a pu
coloniser la région calédonienne à partir du
plateau continental australien ou via la ride de
Norfolk à l'exemple du modèle envisagé par
Bouchet et Poppe (1988) pour le gastéropode

Volutidae Alcithoe. La dispersion étant favorisée
au Néogène alors que l’ensemble Australie/Nou¬
velle-Zélande/Nouvelle-Calédonie était plus rap¬
proché et les rides tectoniques plus prononcées.
Notons qu 'Eumitra n’est connue en Nouvelle-
Zélande que dans le Miocène inférieur (Pakau-
rangi Point, Miocène inférieur) mais serait
présent dans le sud de l’Australie jusqu'au
Pliocène inférieur.

La répartition spatio-temporelle disjointe de
Charitodoron est certainement, avant tout, un
problème de lacune. Mais, on peut penser que
sa distribution actuelle, restreinte, correspond à
une dernière étape dans le processus expan¬
sion/réduction/disparition d’un taxon. Chez les
Volulacea on connaît plusieurs exemples ana¬
logues, j’en exposerai deux :

les Athletinae (Volutidae) ont au Paléogène
une vaste répartition téthysienne et sont abon¬
dantes dans la zone infralittorale. Actuellement
cette sous-famille a une distribution en « taches »
et les 15 espèces vivent essentiellement dans le
bathyal supérieur (moyenne des profondeurs rele¬
vées pour l’Afrique du Sud, 325 m ; l'Australie,
348 m). La principale « tache » se situe le long
des côtes d'Afrique du Sud (11 espèces, Liltved
& Millard, 1986), une espèce vit en Tanzanie,
une autre dans le sud de la Somalie (Rehder,
1981), et deux espèces sont connues du nord-
est de l’Australie (Darragh, 1979) (voir carte
Fig. 25);

le genre Loxotaphrus (Cancellariidae ; Beu &
Maxwell, 1987) est connu de l’Oligocène au
Miocène en Europe et en Inde, du Miocène sud-
australien, et actuellement une seule espèce sub¬
siste dans la province ouest-africaine : Loxota¬
phrus deshayesi (Duval, 1841). On constate
d’autre part que Loxotaphrus possède jusqu’à
l’Oligocène supérieur (observ. inédite) une proto-
conque de type planctotrophe facilitant donc la
dispersion. La perte de la planctotrophie inter¬
vient au cours du Miocène, L. varie férus (Tate,
1888) d'Australie et L. deshayesi sont non-planc-
totrophes (Beu & Maxwell, 1987).

Ces différents exemples de disjonctions d’aires
de répartition évoqués chez les Volutacea ne
permettent pas d'activer les grands problèmes de

Source : MNHN, Paris

MOLLUSCA GASTROPODA : EU MIT RA ET CH A R/TODORON

217

Fig. 25. Carte montrant des disjonctions spatio-temporelles de répartition chez quelques Volutacea :
Volutidae (Athletinac), distribution au Paléogène (a) et actuelle.

Milridae ( Cltaritodoron ), especes paléogènes (c) et distribution actuelle.

Mitridac (Eumitra), espèces fossiles miocènes (e) et distribution actuelle.

Cancellariidae ( Loxolaphrus ), distribution au Miocène (l) et actuelle.

la biogéographie mobiliste. En fait, la distribu¬
tion actuelle de la majorité des organismes
benthiques littoraux (voire bathyaux) et péla¬
giques des zones tropicales et subtropicales s’ex¬
plique souvent au regard de deux événements
majeurs, qui amorcent le dernier cycle de l’évolu¬
tion de la biosphère marine (Spokl, 1983 ; Roux,
1982) :

— la crise climatique à la limite Eocène/Oligo-
cène enregistrée par la plupart des organismes
(Cavelier, 1979), vraisemblablement liée au
déplacement du continent Antarctique (? conjugé
à une « catastrophe cosmique », Raup & Sep-
koski, 1984). La planète rentre alors dans un
cycle glaciaire (Chamley, 1984);

l'interruption de la liaison tropicale mon¬
diale, à la limite Oligocène/Miocène suite à la

jonction des plaques arabique et africaine. Elle
entraîne notamment l'individualisation de deux
vastes provinces tropicales (Indo-Ouest-Pacifique
et Euro-Ouest-Africaine, Lozouet, 1986) et l’in¬
version du sens de la dispersion planctonique
(Spoel, 1983).

En dehors de ces événements (sans négliger
l’importance d’épisodes plus récents tel l’émer¬
sion du seuil de Panama), l’interaction entre les
propriétés biologiques propres à chaque espèce
et les facteurs physico-chimiques, suffit à rendre
compte de la distribution des organismes. C'est
pourquoi la plupart des disjonctions spatio-tem¬
porelles, dans le domaine marin tropical et
subtropical, ne nécessitent généralement pas une
explication fondée sur une paléogéographie anté-
Eocène, ainsi que le souligne Bouchet (1987).

REMERCIEMENTS

Je remercie les différentes personnes qui m’ont
communiqué le matériel sur lequel est fondé ce
travail : P. Bouchet (mnhn). R. Janssen (smf),
R. Kii.burn (nm), C. Levi (mnhn), J. Prudhomme
(Muséum de Bordeaux), B. Richer de Forges
(Orstom, Nouméa). Les critiques et commentaires

de P. Bouchet, W. Cernohorsky (Auckland),
A. Crosnier (Orstom. Paris) et P. Maxwell
(Waimate, Nouvelle-Zélande) m'ont été particu¬
lièrement utiles. A. Waren a préparé la radula
d 'Eumitra.

Source : MNHN, Paris

218

PIERRE LOZOUET

Fig. 26-35. Genre Charitodorun. 26-28. C. barbara, Afrique du Sud, Agulhas Bank (NM b3109), h = 23 mm. - 29-30
et 35, C. agulhasensis, Afrique du Sud, Agulhas Bank (nm a4054), h = 25 mm. - 31-32. C. cancellalus , Oligocène
supérieur. St-Etienne-d'Orthe (mnhn), h = 13.7 mm. 33-34, C. tauzini , Oligocène supérieur. St-Etienne-d'Orlhe,
holotypc (ipm r53074), h = 18,3 mm. — Fig. 26-27 et 30-35 avec placage de chlorure d'ammonium.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : EUM1TRA ET CH A R1TODO RO N

219

Fig. 36-42. 36-38, délai! des premiers tours de spire de Chariiodoron : 36, C. agulhasensis. Afrique du Sud, Agulhas Bank

(nm a4054). 37, C. barbara, Afrique du Sud. Agulhas Bank (nm b3109). 38. C. lauzini sp. nov.. holotype. 39-

40. Fusus scrobiculatus, Oligçcène supérieur d'Allemagne (smf 250349/1). h = 15 mm. 41-42, Parvisipho terebralis
(espèce-type de Parvisipho), Éocène moyen du bassin de Paris, h = 9 mm. Toutes les figures avec placage de chlorure
d'ammonium.

Source : MNHN, Paris

220

PIERRE LOZOUET

Fig. 43-50. Eumitra de Nouvelle-Calédonie. 43-46, Eumitra caledonica : 43-45, holotype. Biocal, stn 61, h = 35 mm
46, paratype, détails premiers tours. Biocal, stn 70, Échelle = 5 mm. 47-50. Eumitra apheles : 47^49, holotype
Musorstom 4. stn 159. h = 17 mm ; 50. paratype, Musorstom 4. stn 159, il = 17 mm. Fig. 44-46, 48-50 avec
placage de chlorure d'ammonium.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : EU MITRA ET CHAR1TODORON

221

Fig. 51-58. Eumitra de la région néo-calédonienne. 51-55. Eumitra richeri, mer du Corail. SW de Mellish Reef.
Corail 2. stn 172 : 51. paratype. il = 19.7 mm ; 52-54. holotype. il = 22.5 mm : 55. paratype. il = 17.6 mm. 56-
58. Eumitra imbricata, mer du Corail. Lansdowne-Fairway, Corail 2, stn 14. holotype. H = 15.6 mm. Fig. 51-52.
54-55, 58 avec placage de chlorure d'ammonium.

Source : MNHN, Paris

222

PIERRE LOZOUET

RÉFÉRENCES BIBLIOGRAPHIQUES

Barnard, K. H., 1959. Contributions to the
knowledge of South African marine mollusca.
Part II. Gastropoda : Prosobranchia Rhachiglossa.
Ann. S. Afr. Mus.. 45 : 1-237.

Barnard, K. H., 1960. The genus Charitodoron
Tomlin, 1932. J. Conch.. 24 (11) : 402.

Beu. A. C., & Maxwell. P. A.. 1987. — A Révision
of the Fossil and Living Gastropods Related to
Plesiolriton Fischer. 1884 (Family Cancellariidae,
Subfamily Plesiotritoninae n. subfam.). N. Z. Geol.
Surv. Pal. Bull., 54 : 1-140.

Bouchet, P.. 1987. — La protoconque des Gastéro¬
podes. Aspects biologiques, taxonomiques et évo¬
lutifs. Thèse de Doctorat d'État (inédite), Paris,

181 p.

Bouchet, P., & Poppe, G., 1988. Deep Water
Volutes from the New Caledonian Région, with a
discussion on Biogeography, Venus, 47 (1) : 15-32.
Cavelier, C., 1979. — La limite Éocène-Oligocène
en Europe occidentale. Mém. Sci. Géol., Strasbourg,
54 : 1-280.

Cernohorsky, W. O., 1970. — Systematics of the
families Mitridae & Volutomitridae (Mollusca :
Gastropoda). Bull. Auckland Inst. Mus., 8 : 1-
190.

Cernohorsky, W. O., 1976. The Mitridae of the
World. Indo-Pacific Mollusca, 3 (17), 273-528.
Chamley, H., 1984. — Les Paléoenvironnements à
la lumière des données océanologiques modernes.
Bull. Soc. géol. Fr., (7) 26 (3) : 407-416.
Darragh, T. A., 1979. — New species of Athleta
(Ternivoluta) and Nannamoria (Mollusca : Vo-
lutidae) from the Capricorn Channel, Central
Queensland, Australia. J. Malac. Soc. Aust., 4 (3) :
129-134.

Finlay, H. J., 1926. A Further Commentary on
New Zealand Molluscan Systematics. Trans. Proc.
N. Z. Inst., 57 : 320-485.

JANSSEN, R., 1979. — Die Mollusken der oberoligozân
(Chattium) im Nordsee-Becken. 2. Neogastropoda.
Euthyneura, Cephalopoda. Arch. Moll., 109 (4/6) :
277-376.

Hornes, M., 1956. — Die fossilen Mollusken des
Tertiàrbeckens von Wien. Univalen. Abhandl. Geol.
R. Anst., 3 : 1-736.

Le Renard, J.. 1989. Clefs de détermination des
petites espèces de Gastéropodes de l'Éocène du
Bassin de Paris. XXXVIII Le genre Parvisipho
Cossmann, 1889. Cah. Naturalistes, 45 (3) : 49-67.
Liltved, B.. & Millar. V., 1986. - Volutidae of
South Africa. The Strandloper, 215 : 2-3.
LOZOUET, P., 1986. Les Gastéropodes Proso-
branches de l'Oligocène supérieur du bassin de
l'Adour (Systématique. Paléoenvironnements, Paléo

climatologie. Palcobiogéographie). Mémoire E.P.H.E.
(Paris) : 475 p.

Marshall, P.. 1918. The Tertiary Molluscan
Fauna of Pakaurangi Point, Kaipara Harbour
Trans. Proc. N. Z. Inst.. 50 : 263-278.

Ponder, W.. & Waren, A., 1988. — Classification of
the Caenogastropoda and Heterogastropoda, a list
of the family-group names and higher taxa. Malac.
Rev., suppl. 4 : 286-324.

Powell, A. W.. & Bartrum, J. A.. 1929. The
Tertiary (Waitematan) Molluscan fauna of Oneroa,
Waiheke Island. Trans. Proc. N. Z. Inst.. 60 (3) •
395-447.

Raup, M., & Sepkoski, J. J., 1984. Periodicity of
extinctions in the géologie past. Proc. Nain. Acad.
Sci. USA. 81 : 801-805.

Rehder, H. A.. 1981. A new species of Volutocorbis
(Volutidae) from Somalia. Nautilus, 95 (4) : 169-
170.

Rossi Ronchetti, C., 1955. — 1 Tipa délia «Concho-
logia Fossile Subapennina » di G. Brocchi. II.
Gastropodi, Scafopodi. Riv. ital. Paleont. Stratigr.
5 (2) : 91-343.

Roux, M., 1982. — De la biogéographie histo¬
rique des océans aux reconstitutions paléobiogéo¬
graphiques : tendances et problèmes illustrés par
des exemples pris chez les Echinodermes bathyaux
et abyssaux. Bull. Soc. géol. Fr., (7) 25 (5-6) : 907-
916.

Saubade, A. M., 1969. Découverte d'une nouvelle
espèce de gastropode dans les faluns de Saint-
Étienne-d'Orthe (Landes). Bull. Soc. Borda (Dax) :
1-3.

Spoel van der. S., 1983. Patterns in the Plankton
Distribution and the Relation to Spéciation : The
Dawn of Pélagie Biogeography. In : R. W. Sims,
J. H. Price & P. E. S. Whaley (eds), « Evolution,
Time and Space : I'he Emergence of the Biosphère ».
Systematics Association. 23 : 291-334, Academie
Press, London.

Tate, R.. 1889. The Gastropods of the Older
Tertiary of Australia. Part II. Trans. R. Soc. Austr.,
11 : 116-174.

Tenison-Woods, 1880. — On some Tertiary Fossils.
Proc. Linn. Soc. N. S. W. , 4 (1) : 1-20.

Thiele, J., 1925. Gastropoda der Deutschen
Tiefsee-Expedition, 2. IViss. Ergebn. dt. Tiefsee-
Exped. " Valdivia ", 17 : 35-282.

Tomlin, J. R., 1932. 9. Reports on the Marine

Mollusca in the Collections of the South African
Muséum, VI-VIII. Ann. S. Afr. Mus.. 30 (2) : 157-
169.

Tomlin, J. R., 1943. New species of Charitodoron.
J. Conch.. 22 (3) : 50.

Source : MNHN, Paris

LTATS DES CAMPAGNES MUSORSTOM. VOLUME 7 RÉSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÊSULT/

9

Mollusca Gastropoda : The Typhinae (Muricidae)
from the New Caledonian région with description
of five new species

Roland HOUART

St Jobsstraat, 8
B-3400 Landen
Ezcmaal. Belgique

ABSTRACT

The New Caledonian species of Typhinae are revised. A total
of 11 species are rccordcd ; 5, ail front deep-sea, are new :
Siphonochelus (S.) angustus: S. (S.) boucheti : S. (S.)
sahaniis : S. (S.) unicornis and S. (? Siphonochelus) undu-
laïus. Ail the species are described and illustrated together

with comparative material. The radulae of 3 species are
illustrated Typhis (Typhina) carolinae Houart, 1987 ;
Siphonochelus fS.) boucheti sp. nov. and S. (S.) sahaniis
sp. nov. Position and angle of anal tubes are considered to
be a good criterion for the séparation of species.

RÉSUMÉ

Mollusca Gastropoda : Les Typhinae (Muricidae) de la
région néo-calédonienne. Description de cinq espèces nouvelles.

L'étude des Typhinae récoltés au cours des différentes
campagnes entreprises depuis 1978 en Nouvelle-Calédonie
a permis de recenser 11 espèces, dont 5 nouvelles, toutes
d’eau profonde : Siphonochelus (S.) angustus : S. (S.)
boucheti ; S. (S.) saltantis: S. (S.) unicornis et S. (?Sipho-
noclielus) undulatus. Les 11 espèces sont décrites, comparées
et illustrées. Les figures reprennent également quelques
espèces comparées provenant d'autres régions du Pacifique.

notamment du Japon et d'Australie. La radula de 3 espèces
est illustrée : Typhis (Typhina) carolinae Houart, 1987 ;
Siphonochelus (S.) boucheti sp. nov. et S. (S.) sahaniis sp.
nov. La position des tubes aperturaux (canal anal) et l’angle
qu'ils forment sont considérés comme de bons critères pour
la séparation des espèces. Par ailleurs d'autres points ont été
retenus et ont servi à la comparaison et à la séparation
d'espèces : forme et taille de la protoconque, sculpture et
morphologie des varices, sculpture axiale intervaricale et
sculpture spirale.

Houart, R.. 1991. Mollusca Gastropoda : The Typhinae (Muricidae) from the New Caledonian région with description of
five new species. In : A. Crosnier & P. Bouchet (eds). Résultats des Campagnes Musorstom. Volume 7. Mc ni Mus. nain. H ht. nat.. (A). ISO
223-241. Paris ISBN : 2-85653-180-6.

Publié le 20 mars 1991.

Source : MNHN, Paris

224

ROLAND HOUART

INTRODUCTION

Several French expéditions hâve collected a
considérable amount of marine material from
deep water in the New Caledonian région,
including many species of Muricidae. Among
that material it has been possible to select
an important collection of Typhinae, including
many unknown species. Other Muricidae from
the New Caledonian région are being studied by
the author (in press) or hâve already been
reported in other papers (Houart, 1983, 1986,
1987a, 1987b, 1990).

The subfamily Typhinae is well studied and
several authors (Keen, 1944; Vella, 1961;
Gertman, 1969: d'Attilio, 1975, 1976, 1979
and d'Attilio & Hertz, 1988) hâve done much
for a better understanding of this group and its
classification. The classification and methods
used in the présent paper are based mainly on
the conclusions of these previous authors, but
also on personal observations.

In their recent paper, d'Attilio & Hertz
(1988) reinstate the family Typhidae Cossmann,
1903, which they divide into two subfamilies :
Typhinae and Tripterotyphinae. The purpose of
this paper being not the discussion of the
supraspecific classification of Typhinae, but only
the review of the New Caledonian species, no
other remarks will be given here on that study.

The Typhinae include a total of 45 Recent and
77 fossil species. No records were known from
the New Caledonian waters before Houart
(1986), except one or two lots in European
Muséums (irsnb i.g. 10591) but to my know¬
ledge nothing has been published about them.
Most Typhinae live in deep water, between 100
and 500 m, with an important radiation around
300-450 m, although some species occur in
shallower waters (in New Caledonia Typhis
carolinae Houart, 1987, and Typhis neocaledo-
nicus Houart, 1987).

The species are characterized by their sraall
size, whitish coloured shell and by the presence
of hollow tubes, situated between each pair of
varices, of which the function is not precisely
known, although it is certain they are anal tubes.
Thèse anal tubes are gradually closed and bro-
ken during the growth of the shell, only the last
tube is open and generally long to very long
when intact. Other constant characteristics are
the closed siphonal canal and the 4 (very rare-
ly 5) varices per whorl. The group of species with
three varices per whorl, formerly considered
Typhinae, and consisting of the généra Pteroty-
phis Jousseaume, 1880, Tripterotyphis Pilsbry
& Lowe, 1932, Prototyphis Ponder, 1972, and
Cinclidoiyphis du Shane, 1969, hâve been transfe-

Fios I. Schcmalic drawing of a Typhinae radula. kt : rachidian tooth : LT : latéral toolh : MD : marginal dcnticle : mc :
marginal cusp ; t.c : latéral cusp : od : outer dcnticle ; ce : central cusp ; ld : latéral dcnticle. Terminology mainlv based
on Fujioka (1985) and Kool (1987).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : TYPHINAE

225

red to the subfamily Muricinae by d’Attilio
(1982). and recently (d’Attilio & Hertz, 1988)
to the subfamily Tripterotyphinae, with the
exception of Cinclidotyphis.

The protoconch of ail known Recent species is
smooth and paucispiral, formed by 1.25 to
2 whorls, indicating non-planktotrophic larval
development, and thus a short to non existent
free swimming larval stage. The operculum has
an apical nucléus.

The radula consists of a rachidian tooth
generally bearing 3 main cusps and 2 latéral
denticles, and a pair of broad and curved latéral

tooth. Latéral denticles of the central tooth are
not always symmetrical, sometimes they are bifid
or divided into 2 or more, smaller denticles. The
presence and placement of the marginal denticles
or cusps are erratic. Aberrant radular features in
Typhinae can be observed in Hausiellotyphis
cumingii (Broderip. 1833) or Typhisopsis coro-
natus (Broderip. 1833) from the eastern Pacific
région (Radwin & d'Attilio, 1976 : 195. 213).
Otherwise stated. the material is housed in the
Muséum national d'Histoire Naturelle, Paris
(mnhn).

MATERIAL AND METHODS

The material was collected during the fol-
lowing cruises (see Richer de Forges, 1990,
for additional data) :

(1) Aboard R. V. “ Vauban ”, off South New
Caledonia, by P. Bouchet and A. Waren (1978-
79).

(2) Programme Lagon, aboard R. V. "Van-
ban ”, conducted by B. Richer de Forges (1984-
1989).

(3) Biocal cruise, aboard R. V. " Jean Char¬
cot ”, under the direction of C. Lévi (1985).

(4) Musorstom 4 cruise, aboard R. V. "Vau¬
ban ”, under the direction of B. Richer de
Forges (1985).

(5) Musorstom 5 cruise, aboard R. V. "Corio-
lis ”, under the direction of B. Richer de Forges
(1986).

(6) Chalcal 2 cruise. aboard R. V. “ Coriolis ”,
under the direction of B. Richer de Forges
(1986).

Abbreviations :

a) preceding station numbers :

DW : Drague Waren (Waren Dredge)

CC : Chalut à crevettes (Shrimp Trawl)

CP : Chalut à perche (Beam Trawl)

DC : Drague Charcot (Charcot Dredge)

Ail stations of programme Lagon are dredgings
using a Charcot dredge.

b) after data :

spm(s) : live-taken specimen(s) présent in
sample

sh(s) : only empty shells présent in sample

The main features here retained for the sépara¬
tion of species are the form and size of the

protoconch, the position, form and angle of the
anal tubes, the sculpture and morphology of the
varices, the intervarical axial sculpture (except
growth fines) and the spiral sculpture.

The following measurements are used (Fig. 2) :
a. Length of the shell : taken from the apex to
the tip of the siphonal canal.

B. Breadth of the shell : largest breadth, not
including anal tubes.

c. Length and breadth of the aperture : largest
measurements, taken from the inner side of
the peristome.

D. Length of the siphonal canal.

E. Length of apertural anal tube.

F. Breadth of the apertural anal tube : largest
breadth, taken near the base.

G. Tube angle : the shell is placed with the apex
above, the aperture facing left ; the angle is
measured between the axis of the shell and
the axis of the anal tube, near its point of exit.

Muséums where material is deposited are :
ams : The Australian Muséum. Sydney.
nmnz : National Muséum of New Zealand,
Wellington.

nsmt : National Science Muséum. Tokyo.
mnhn : Muséum national d’Histoire naturelle.
Paris.

usnm : National Muséum of Natural History.
Washington. D.C.

irsnb : Institut Royal des Sciences Naturelles de
Belgique. Bruxelles.

Source : MNHN, Paris

ROLAND HOUART

226

SYSTF.MATIC ACCOUNT

Family MURICIDAE Rafinesque, 1815
Subfamily TYPHINAE Cossman, 1903
Genus TYPHIS Montfort, 1810
Subgenus TYPHINA Jousseaume, 1880

Type-species by original désignation : Typhis
belcheri Broderip, 1833.

Typhis (Typhina) imperia lis
Keen & Campbell. 1964
Figs 10. 40, 60

Typhis ( Typhina) imperialis Keen & Campbell. 1964 :
46, pl. 8. figs 1-4. — Radwin & d'Aitilio, 1976 :
207. pl. 32, fig. 8.

Material examined. New Caledonia. Biocal :
stn DW 64. 24"48'S, I68°09' E, 250 m. 3 Septcmber
1985 : 2 shs.

Musorstom 4 : stn DW 222, 22“58' S, I67°33' E,
410-440 m. 30 Septcmber 1985 : 1 sh.

Chesterfield Islands. Musorstom 5 : stn DW 274,
24"45'S. 159"41 ' E, 285 m. 9 October 1986 : 1 sh.

Type locality. Trawled oIT Tosa, Japan,
33°20'N, 138°40'E, 200 m.

Description. Shell moderaleiy large and
shouldered. up to 13.5 mm (mnhn, Musorstom
4, stn DW 222). Spire high. consisting of 1.5 pro-
toconch whorls and 5 shouldered teleoconch
whorls. Suture of whorls deeply impressed. Pro-
toconch elongate, somewhat keeled and smooth.
Last whorl bearing 4 foliated varices, ventrally
ornamented with crenulations with strong up-
ward, curved, sealed spine on shoulder. No spiral
sculpture. Axial sculpture consisting only of fine

Source : MNHN, Paris

MOLLUSCA GASTROPODA : TYPHINAE

227

growth striae. A rounded anal tube originales
nearest to preceding varix, forming an angle of
approximately 70" with the axis of the shell.

Aperture roundly ovate, edge erect and smooth,
forming an entire peristome. Siphonal canal long
and broad, sealed, ornamented with a broad
fluted spine near its base. Colour creamy-white
with brown-coloured last anal tube in juvénile
specimens.

Measurements (illustrated specimen). a-b :
13.5 x 7.1 mm. — c : 2.5 x 1.9 mm. — D :
5.3 mm. e : tube broken. - F : 1.1 mm. g :
70".

Discussion. — This species is related to Typhis
montforti A. Adams, 1863, and T. teramaehii
Keen & Campbell, 1964, but the shell differs
from both by the presence of a broad, fluted
spine on the siphonal canal and by the angle of
the anal tube. The présent material represents a
wide geographical range extension for this appa-
rently rare species, previously known only from
the type locality.

Typhis ( Typhina) virginiae Houart. 1986
Figs 12, 41. 62

Typhis (Typhina) virginiae Houart, 1986 : 440, pl. 2,
'fig. 7.

Ma terial cxamined. — New Caledonia. “ Vauban "
1978-79 : stn 2, 22" 17'S. 167° 14' E, 425-430 m. type
locality.

Biocal : stn DW 77, 22°15'S, 167° 15' E. 440 m.
5 September 1985 : 9 spms.

Musorstom 4 : stn DW 226, 22°47' S, 167°22' E,
390 m. 30 September 1985 : 1 sh. Stn CC 246.
22"08'S. 167"11'E, 410-420 m. 3 October 1985 :
5 shs. Stn CC 247, 22°09' S. 167" 13' E. 435-460 m, 4
October 1985 : 2 shs.

Type locality. South of New Caledonia,
22°17'S, 167" 14' E, 425-430 m.

Description. Shell small, délicate, triangu-
lar, up to 7.8 mm (holotype). Spire moderately
high, consisting of 1.25 protoconch whorls and
4 angulate teleoconch whorls. Suture of whorls
slightly appressed. Protoconch smooth and round¬
ed, glossy. Last whorl bearing 4 thin, sharp
varices, ending in an acute and curved open
spine. Apertural varix bearing a winglike flange,
extending from the shoulder spine to approxima¬

tely the 3/4 of the siphonal canal. Spiral sculp¬
ture obsolète. A moderately long anal tube
originates between each pair of varices, situated
slightly nearer to succccding varix and forming
an angle of approximately 25° with the axis of
the shell. Other axial sculpture obsolète except
fine growth striae.

Aperture small and ovate, edge forming an
entire, erect peristome. Siphonal canal modera¬
tely long, sealed and smooth, tapering at the end.
slightly bent to the right. Colour of the shell
creamy white, siphonal canal and upper whorls
pale brown ; a pale brown band also sometimes
apparent at the base and the anterior part of the
anal tubes.

Measurements (holotype). — a-b : 7.8 x
4.1 mm. c : 1.4 x 1.1 mm. d : 2.8 mm. —
E-F : 2.9 x 0.6 mm. G : 25".

Discussion. The species was originally
compared with Typhis (Typhina) pauperis Mes-
tayer, 1916, and Typhis ( Typhina) bivaricata
Verco, 1909. From T. pauperis the shell dilTers
by its lower spire, its lower and more globose
protoconch, the position of the anal tubes and
the smooth and sharp varices. It differs from
T. bivaricata by its larger size, its longer sipho¬
nal canal and its smooth and sharp varices. Both
T. pauperis and T. bivaricata differ by having
curving crenulations on the varices, mostly on
the outer apertural lip.

Typhis (Typhina) carolinae Houart, 1987
Figs 3-4, 11, 42. 61

Typhis ( Typhina) carolinae Houart. 1987 : 204, figs 2-

'4. 12, i3.

Material oxamined. New Caledonia. Lagon :
stn 120. 22°28'S. 166'44' E. 46 m. 23 August 1984 :
1 spm. — Stn 296, 22°41'S. 166’44'E. 26 m.
26 November 1984 : I spm. Stn 326. 22°26'S,
167°02'E, 67 m, 28 November 1984 : 1 spm (ams c
153702). Stn 354. 22"32' S, 167‘W E. 78 m.
29 November 1984 : 1 spm. holotype. Stn 382,
22-33' S. 167" 14' E. 57 m. 22 January 1985 : I spm.

Stn 383. 22"32' S. 167" 13' E. 62 m. 22 January 1985 :
I spm. Stn 403, 22"35' S. 167"18' E, 45 m. 23 January
1985 : 1 spm. Sln 405, 22°38' S. 167"20'E. 27 m.
23 January 1985 : 1 spm. (nmnz me 47730). — Stn 562.
22"44' S. 166°59‘ E. 48 m. 16 Julv 1985 : 1 spm. Stn
572, 22°52' S. 167 00'E. 65 m. 17 July 1985 : 1 spm.
(R. Houart coll.). Stn 598, 22“ 19' S. 167"06’ E. 73-
75 m. 5 August 1986 : 1 spm. Stn 603. 22° 16' S

Source : MNHN, Paris

228

ROLAND HOUART

167"05' E, 78-80 m. 5 Augusl 1986 : 1 spm. Stn 632,
21 ”57' S, 166 "50' E. 44-45 m, 6 August 1986 : I spm.

Type locality. New Caledonia, Grand
Récif Sud, Lagon. 22“32' S, 167°02'E, 78 m.

Description. Shell large and spinose, up to
20.5 mm (holotype). Spire moderately high,
consisting of 1.5 protoconch whorls and 5 shoul-
dered, spinose, teleoconth whorls. Suture of
whorls impressed. Protoconch rounded, smooth
and glossy. Last whorl bearing 4 spinose and
ventrally squamous varices, ornamented with
small, backwardly curved. open spinelets. Shoul-
der spine long and strongly upwards curved.
Spiral sculpture consisting of 3 to 4 very shallow,
sometimes squamous, low cords on the posterior
part of last whorl. Axial sculpture of fine growth
striae. A rounded anal tube originates about
midway belween each pair of varices, slightly
nearer preceding varix, forming an angle of 60 to
90° with the axis of the shell. Last tube long to
very long and hollow, others gradually shorter
and closed.

Aperture roundly-ovate to rounded, edge erect
and smooth, forming an entire peristome. Sipho-
nal canal sealed, long to very long, posteriorly
fiat and broad with spinelike projections poste¬
riorly. Colour creamy-white to light brown,
shoulder darker. Operculum with apical nucléus.
Radula : rachidian bearing 3 cusps and 2 latéral
denticles. Central and latéral cusps of approxi-
mately the same size ; latéral denticles slightly
shorter and narrower ; 1 or 2 small outer
denticles may erratically be présent on the base
of central cusp.

Measurements (holotype). a-b : 20.5 x
11.2 mm. c : 4 x 3.5 mm. D : 10.5 mm.

E-F : 13.3 x |.5 mm. g : 90°.

Discussion. This species was originally com-
pared with Tvphis (Typhina) montforti A. Adams,

Figs 3-4. Radula of T. carolinae Houart, 1987 (scalc Unes
10 |am).

1863, T. (T.) yatesi Crosse & Fischer, 1865,
T. (T.) imperialis Keen & Campbell, 1964 and
T. (T.) teramachii Keen & Campbell. 1964.
The shell dififers from these four species in having
more spinose varices ; the varical ornamentation
being dorsally bent, while the other species show
ventrally bent crenulations. Other différences
with relaled species are the position and angle of
anal tubes ; ornamentation of the siphonal
canql ; form of the protoconch and sculpture of
the shell.

Subgenus TALITYPHIS Jousseaume, 1882

Type-species by original désignation : Typhis
expansus Sowerby. 1874.

Typhis (Talityphis) neocaledonicus Houart, 1987
Figs 13, 44, 66

Typhis (Talityphis) neocaledonicus Houart, 1987 : 208.
figs 8. 9, 16.

Material examined. New Caledonia. Lagon •
stn 416, 22"38'S, 167° 14' E, 40-50 m : 1 spm,
holotype.

Type locai.ity. New Caledonia, Grand
Récif Sud, Lagon, 22°38’ S, I67°14' E, 40-50 m.

Source : MNHN, Paris

MOLLUSCA OASTROPODA : TYPHINAE

229

Description. Shell relatively small for the
subgenus and broadly fusiform, up to 18 mm.
Spire moderately high, consisting of 1.5 proto-
conch whorls and 5 shouldercd and angulate
teleoconch whorls. Suture of whorls impressed.
Protoconch rounded, somewhat flattened and
smooth. Last whorl bearing 4 laminate and
sharp varices. Last varix broad and expended,
extending from the uppcrmost part of the shoul-
der spine to almost the tip of the siphonal canal.
The partition is well developed. Varices ending in
a sharp open shoulder spine. No apparent spiral
sculpture but very shallow, low cords on poste-
rior side of varices. A rounded anal tube origi-
nates at the shoulder margin, just in front of
preceding varix, forming an angle of 70” with the
axis of the shell. Last tube hollow and long,
others short and closed.

Aperture rounded, edge erect and smooth,
forming an entire peristome. Siphonal canal
moderately long, broad and sealed, strongly bent

backwards on its tip. Shell colour pale brownish
with some brown spots on the edge of the
aperture and on the siphonal canal.

Measurements (holotype). a-b : 18 x
13 mm. — c : 3.5 x 2.9 mm. — d : 6.5 mm. —
e-f : 9.4 x 1.9. g : 70°.

Discussion. T. neocaledonicus may be
compared with Typhis (Talityphis) bengalensis
(Radwin & d’Attilio, 1976) from the Bay of
Bengal, but that species has a much smaller shell
(7.6 to 9.5 mm), with no spiral sculpture, a larger
aperture and a different arrangement of the anal
tubes, situatcd approximately medially between
each pair of varices. T. (T.) campbelli (Radwin
& d’Attilio. 1976), has a more slender and higher
protoconch ; the shoulder spines are not recur-
ved and the position of the anal tubes is
different, also situated medially between each
pair of varices.

Genus SIPHONOCHELUS Jousseaume, 1880
Subgenus SIPHONOCHELUS Jousseaume. 1880

Type-species by original désignation : Typhis
arcuatus Hinds, 1843.

Siphonochelus (Siplionochelus) pavlova
(Iredale. 1936)

Figs 14, 35. 45, 56, 63

Typhina pavlova Iredale, 1936 : 324, pl. 24, fig. 12. —
Radwin & d'Attilio, 1976 : 205, pl. 31, fig. 6. —
Kaicher, 1978 : 1563. — Houart. 1986 : 435, pl. 3,
figs 8, 8A.

Material examined. — New Caledonia. “ Vauban "
1978-79 : stn 40, 22"30' S, 166"24' E. 250-350 m, 7 June
1979 : 10 shs.

Biocal : stn DW 77, 22°15'S, 167°15' E, 440 m.
5 September 1985 : 1 spm.

Musorstom 4 : stn CC 246. 22°08' S. 167° 11' E. 410-
420 m, 3 October 1985 : 1 sh. - Stn CC 247. 22°09' S,
167°13' E. 435-460 m, 4 October 1985 : 1 spm.

Chesterfield lslands. Musorstom 5 : stn DW 301,
22“07'S. 159 Ü 25'E. 487-610 m : 1 spm.

Australia. Trawled olT Cape Moreton. South
Queensland : 2 spms (R. Houart coll.).

Type locality. — Australia. east of Sydney,
200 m.

Description. Shell small, biconic. up to
10 mm (mnhn, Musorstom 4. stn CC 247). but
larger (up to 16 mm) in Australian specimens.
Spire high, consisting of 1.5 to 1.6 protoconch
whorls and 5 somewhat shouldered teleoconch
whorls. Suture of whorls impressed. Protoconch
rounded. elongate and smooth. Last whorl bea¬
ring 4 rounded varices, joined to varix of
preceding whorl by a butlress. No spiral sculp¬
ture. Axial sculpture consisting of a very shallow
axial ridge and fine growth striae. A long,
somewhat curved, tubular anal tube originates
from the succeeding varix, forming an angle of
approximately 40 to 45° with the axis of the shell.
Tube of apertural varix long and hollow. others
short (broken) and closed.

Aperture roundly-ovate, edge erect and smooth,
forming an entire peristome. Siphonal canal
long, sealed and smooth. slightly bent to the

Source : MNHN, Paris

230

ROLAND HOUART

right and tapering on its tip. Shell whitish to
light brown with brown maculations on the base
of the siphonal canal and on the buttresses on
the suture.

Measurements (illustrated specimen, mnhn).
a-b : 10 x 4.9 mm. — c : 1.7 x 1.4 mm. d :
3.5 mm. — e-f : 2.9 x 9 mm. — g : 40 to 45°.

Discussion. — For the différences with Sipho-
nochelus (S.) sallantis sp. nov. see that species.
There are no noticeable différences with the
Australian shells, except these are somewhat
more globose with slightly more rounded varices.

Known in Australia from the coast of New
South Wales (Iredale, 1936) and from Cape
Moreton, South Queensland (Radwin & d’Atti-
lio, 1976 : PI. 31, Fig. 6 and R. Houart coll.).

Siphonochelus (Siphonochelus) angustus sp. nov.

Figs 15-16, 43, 65

Material examined. Chesterfield Islands. Mus-
ORSTOM 5, stn DW 304, 22“10' S, 159°26' E, 385-420 m :
2 shs.

Type material. Holotype mnhn, Musor-
tom 5, stn DW 304 (6.7 x 3.4 mm) ; 1 para-
type mnhn, same station (6.3 x 3.3 mm).

Type locality. Coral Sea, Nova Sea-
mount, Musorstom 5, 22°10'S, 159°26'E, 385-
420 m.

Description. Shell small and fusiform,
biconic, up to 6.7 mm (holotype). Spire high,
consisting of 1.5 protoconch whorls and 4 con-
vex teleoconch whorls. Suture of whorls impres-
sed when visible. Protoconch broad, rounded,
slightly flattened and glossy. Last whorl bearing
4 rounded varices. No spiral sculpture. Axial
sculpture consisting of one, sometimes shallow
or obsolète, axial ridge, situated midway be-
tween each pair of varices. Shell slightly exca-
vated between axial ridge and succeeding varix.
A flattened and broad anal tube originates from
succeeding varix, forming an angle of approxi-
mately 10 to 15° with the axis of the shell ; its
broad and flattened base masks the suture of
whorls. Other axial sculpture of fine growth
striae.

Aperture ovale and smooth. partially broken.

Siphonal canal short, broad at the base and
narrower at its tip, smooth, tubular and sealed.
Shell ivory white and glossy.

Measurements (paratype). a-b : 6.3 x
3.3 mm. c : 1.4 x 1.0 mm. d : 1.7 mm.

e : tube broken. F : 1.2 mm. G : 10 to 15°.

Discussion. The shell of Siphonochelus
japonicus (A. Adams, 1863) is brown coloured,
has more pronounced axial ribs and is deeply
excavated between each varix and axial rib ; the
suture is deeper and the anal tubes are more
detached from the whorls ; the varices are
weaker and the protoconch is higher and more
rounded.

? Siphonochelus svringianus (Hedley, 1903) from
shallower water has a brown coloured shell, it is
barely half the size for the same number of
teleoconch whorls ; its protoconch is slightly
smaller and non acute ; its intervarical costae are
more apparent ; the teleoconch whorls are more
detached from each other and the shoulder is
stronger and broader. The varices and anal tubes
are more numerous for S. syringianus : 6 on
first teleoconch whorl and 5 from second teleo¬
conch whorl onwards, including the last whorl
while S. angusius has 4 varices and 4 anal tubes
on each whorl.

Siphonochelus boucheti sp. nov. has a compa-
ratively larger shell, with a more rounded proto¬
conch, smaller anal tubes and broader shell. It
has more pronounced and erect axial ridges and
varices.

Etymology. — Named angusius due to its
relatively narrow shell.

Siphonochelus (Siphonochelus) boucheti sp. nov.

Figs 5-6, 17-18, 47. 68

Material examined. New Caledonia. Biocal :
stn DW 46, 22"53' S, \bT\T E. 570-610 m, 30 August
1985 : 2 spms. Stn DW 51,23”05' S, 16745' E, 680-
700 m, 31 August 1985 : 6 spms. — Stn CP 57
23‘44' S, 166"58' E. 1 490-1 620 m. 1 September 1985 ■
1 sh. Stn DW 66, 24°55’ S, 168"22' E, 505-515 m,
3 September 1985 : 20 spms. — Stn DW 70, 23"25' S,
167 0 53'E, 965 m, 4 September 1985 : 1 sh.

Chalcal 2 : stn DW 72. 24"55' S, 168°22' E. 527 m.
28 October 1986 : 21 spms.

Chesterfield Islands. Musorstom 5 : stn DC 358
19°39'S, 158°47'E, 680-700 m, 18 October 1986 :
1 spm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : TYPHINAE

231

Figs 5-6. Radula of S . haucheti sp. nov. (scale lines
10 |xm).

Type material. Holotype mnhn, Chalcal
2, stn DW 72, 10.1 x 5 mm ; 20 paratypes, same
station (mniin, nmnz, ams. usnm, nsmt. Natal
Mus., R. Houart coll.).

Type locality. South of New Caledonia,
Chalcal 2, stn DW 72, 24°55'S, 168°22'E,
527 m.

Description. Shell small and fusiform,
biconic, up to 10.5 mm (specimen from Biocal,
stn DW 66). Spire high, consisting of 1.6 pro-
toconch whorls and 5 shouldered teleoconch
whorls. Suture of whorls impressed. Protoconch
rounded, smooth and glossy. Last whorl bearing
4 rounded varices, angulate at the shoulder. No
spiral sculpture. Intervarical axial sculpture con¬
sisting of a strong rounded ridge, situated about
midway between each pair of varices, slightly
nearer to succeeding varix. Shell excavated be¬
tween axial ridge and succeeding varix. A flat-
tened and arched anal tube originates from the

succeeding varix ; broad at the base and gra-
dually narrower at its tip, strongly upward bent,
forming an angle of approximately 20 to 25°
with the axis of the shell. Other axial sculpture of
fine growth striae.

Aperture ovate, edge erect and smooth, form¬
ing an entire peristome. Siphonal canal short,
broad at the base and narrower at its tip.
smooth, tubular and sealed. Shell ivory white
and glossy.

Radula : rachidian bearing 5 cusps and 2 la¬
téral denticles. Laterals cusps rarely bearing
outer denticles. Latéral denticles erratically di-
vided into 2 or 3 smaller denticles.

Measurements (holotype) : a-b : 10.1 x
5 mm. - c : 2.5 x 1.7 mm. d : 2.6 mm.
e-f : 2.5 x 1.2 mm. G : 20 to 25°.

Discussion. S. boucheti is related to Sipho-
nochelus nipponensis Keen & Campbell, 1964 but
the shell has more upward vertically bent anal
tubes which are flatter on the base : they origi-
nate immediately from the varix in S. nipponensis
while they are somewhat séparated in S. bou¬
cheti. The varices are more strongly shouldered
than in S. nipponensis ; the protoconch of S.
boucheti is approximately 50 % larger, and the
aperture is narrower.

Siphonochelus japonicus (A. Adams, 1863), a
smaller related species. has been compared with
adult specimens of S. boucheti and also with an
approximately equal-sized juvénile. S. boucheti
has a translucent white shell while S. japonicus
has a brownish to dark brown shell, but other
différences are the more shouldered whorls in S.
boucheti , which also has a more globose and
almost twice larger protoconch ; the varices are
strongly shouldered and the anal tubes are
slightly joined to them while these of S. japoni¬
cus are rounded with the anal tubes distinctly
joined ; these anal tubes being flatter in S.
japonicus.

Siphonochelus generosus Iredale, 1936. a South-
eastern Australian species is also similar to the
new species but S. boucheti has a twice larger
and more globose protoconch, more fusiform
whorls, more erect apertural lip and slightly
more upwardly bent anal tubes.

Siphonochelus solus Vella, 1961, is a more
fusiform species with smaller protoconch and

Source : MNHN, Paris

232

ROLAND HOUART

weaker axial costae. The new species also has
slightly more upwardly bent anal tubes and the
axial costae are more separated from the anal
tubes.

5. tillierae Houart, 1986, is a narrower and
comparatively smaller shell. The anal tubes do
not originate from the varices and are rounded
and straight, while flattened and arched for the
new species. The axial ridge is also stronger and
more évident in S. boucheti.

Etymology. Named in honour of Dr
P. Bouchf.t (mnhn), who shared in most of
the recent French expéditions in New Caledonia.

Siphonochelus (Siphonochelus) suit antis sp. nov.

Figs 7-8, 9. 19-20, 46, 70

Material examined. New Caledonia. Biocal :
stn DW 64, 24°48'S, 168°09' E, 250 m, 3 Septembcr
1985 : 5 spms.

Chesterfield Islands. Musorstom 5 : stn DW 263.
25°2T S, 159”46' E, 150-225 m. 8 October 1986 :
9 spms. Stn DW 274. 24"45' S. 15941'E. 285 m,
9 October 1986 : 1 spm. - Stn DW 303, 22°12' S.
159°23' E. 332 m. 12 October 1986 : 1 spm. Stn DW
304, 22“10' S. 159’26' E, 385-420 m. 12 October 1986 :
1 sh.

Type material. - Holotype mnhn, Musor-
stom 5, stn 303. 9.2 x 4.1 ; 1 paratype mnhn,
Musorstom 5, stn 304.

Type locality. Coral Sea, Nova Sea-
mount. Musorstom 5, stn DW 303, 22°12'S,
159°23' E, 332 m.

Description. — Shell small and fusiform,
biconic, up to 9.2 mm (holotype). Spire high,
consisting of 1.5 to 1.75 protoconch whorls and
4 slightly angulate and shouldered teleoconch
whorls. Suture of whorls impressed. Protoconch
small, rounded and glossy. No spiral sculpture.
Axial sculpture consisting of 4 sharp varices and
fine growth striae. A long, fused. slightly angu¬
late and slender anal tube originates from the
varices, forming an angle of approximately 25 to
30° with the axis of the shell. Tube of apertural
varix very long and hollow, others broken and
closed.

Aperture rounded and very small, edge erect
and smoolh, forming an entire peristome. Sipho-
nal canal long and fused, broad at the base.

gradually narrower and tappering at its tip, bent
to the right. Shell glossy white, often with more
or less visible brown maculations at the base of
the anal tubes, siphonal canal and remnants of
previous siphonal canals.

Radula : rachidian bearing 3 cusps, 2 small
latéral denticles and 2 small marginal den-
ticles. Latéral denticles erratically divided into
2 smaller denticles.

Mf.asuremknts (holotype). a-b : 9.2 x
4.1 mm. c : 1.4 x 1.0 mm. — d : 3.6 mm. —
E-F : 3.8 x 0.7 mm. g : 25 to 30".

Figs 7-8. — Radula of S. saltaniis sp. nov. (scalc lines
10 nm).

Discussion. Compared with S. pavlova
(Iredale, 1936), the shell is twice smaller for the
same number of teleoconch whorls ; the aperture
is also smaller. The protoconch is comparatively
smaller, more rounded while that of S. pavlova is
large and elongate. S. pavlova has a more
globose shell with more rounded varices and
more apparent buttresses where the varices join

Source : MNHN, Paris

MOLLUSCA GASTROPODA : TYPH1NAE

233

the preceding whorl. and with a shallow axial
ridge.

Typhis (Typhina) virginiae Houart, 1986, an-
other recenlly named species from New Caledo-
nia has a more angulate shell with a larger
aperture ; the axial lamellae are separated from
the anal tubes and are sharper while the anal
tubes are less backwards bent and more horizon¬
tal. The suture is more impressed in T. virginiae
and the buttresses formed where the varices or
axial lamellae join the preceding whorl are
shallow while they are large and very apparent in
S. saltantis.

Fig. 9. Opcrculum of S. sait amis sp. nov. (scalc line
0.5 mm).

Etymology. Saltantis = dancer. Named
for ils similarity with Siphonochelus pavlova
(Iredale, 1936).

Siphonochelus (Siphonochelus) unie omis sp. nov.
Figs 21-23, 48, 67

Material examined. New Caledonia. Biocal :
sin DW 44. 22'47' S. 167" 14' l£. 440-450 m. 30 August
1985 : 1 spin. Stn DW 46. 22°53' S. 167"17' E. 570-
610 ni, 30 August 1985 : 3 spms.

Musorstom 4 : stn DW 226. 22"47' S. \(sT 22 ' E.
390 m, 30 September 1985 : 1 sh.

Type materiai.. Holotype mnhn, Musor¬

stom 4, stn DW 226, 12 x 5.5 mm ; 1 paratype
mnhn. Biocal, stn DW 44 ; 3 paratypes, Biocal.
stn DW 46 (mnhn. nmnz, R. Houart coll.).

Type locality. New Caledonia. Musor¬

stom 4. stn DW 226, 22°47' S, 167°22' E, 390 m.

Description. Shell small and fusiform.
biconic, up to 12 mm (holotype). Spire high.
consisting of 1.5 to 1.75 protoconch whorls and
5 fusiform and slightly shouldered teleoconch
whorls. Suture of whorls impressed. Protoconch
large, rounded and glossy. No spiral sculpture.
Axial sculpture consisting of 4 more or less sharp
varices and a very shallow axial ridge. situated
nearer to preceding varix. No other axial sculp¬
ture. A long, fragile and almost horizontal anal
tube originates from the apertural varix. forming
an angle of 70 to 75° with the axis of the shell
(paratype mnhn. stn DW 46). Other tubes bro-
ken.

Edge of aperture slightly erect, ovate. forming
an entire peristome. Siphonal canal moderate in
length. broad at the base and narrow at its tip,
smooth. Colour white, with a shallow pale
brown band at the base of the siphonal canal.

Measurements (paratype mnhn). a-b :
9.7 x 4.7 mm. c : i.8 x 1.3 mm. D :
3.3 mm. e-f : 4.5 x 0.8 mm. — G : 75°.

Discussion. — Compared with Siphonochelus
houclieti, S. unicornis has a larger and more
elongate protoconch and a larger aperture. The
axial ribs are weaker and the anal tubes more
backwards bent. being almost horizontal. Sipho¬
nochelus (Siphonochelus) erythrostigma Keen &
Campbell, 1964, has a more rounded and smaller
protoconch, rounded and spirally sculptured
varices, globose and upwardly bent anal tubes.

No other related species is known.

Etymology. — Named unicornis for its long
apertural anal tube.

Siphonochelus (?Siphonochelus) undulutus

sp. nov.

Figs 24-25. 49. 69

Material examined. New Caledonia. Musor¬
stom 4 : stn DW 203, 22‘36' S. I67"05' E. 105-110 m.
27 September 1985 : 1 sh.

Type material. Only known from the
holotype, mnhn.

Type locality. New Caledonia, Musor¬
stom 4, stn DW 203. 22°36' S, 167°05' E, 105-
110 m.

Source : MNHN, Paris

234

ROLAND HOUART

Dkscription. Shell very small, fusiform and
biconic. Spire high, consisting of 1.75 proto-
conch whorls and 4 teleoconth whorls. Suture of
whorls slightly appressed. Protoconch slightly
elongate, convex and smooth. Last whorl bea-
ring 4 rounded varices, extended on previous
whorl by a small buttress. Other axial sculpture
consisting of one elongate ridge, situated nearest
to preceding varix ; shell excavated between axial
ridge and succeeding varix. Spiral sculpture
consisting of 5 cords, most évident on the varices
and axial ridges. A short, rounded anal tube
originates from succeeding varix, forming an
angle of approximately 20 to 25° with the axis of
the shell.

Aperture roundly-ovate, forming an entire
peristome, smooth and slightly erect. Siphonal
canal short, smooth and fused. sealed and ta-
pering on its tip. Colour ivory-white with traces
of pale brown on the anterior part of last whorl.

Measurements (holotype). — a-b : 5.9 x
3.0 mm. — c : 1 x 0.8 mm. — D : 1.4 mm.
e : tube broken. F : 0.6 mm. G : 20 to 25°.

Discussion. S. undulatus is here lentatively
included in S. ( Siphonochelus), its shell sculpture
being somewhat unexpected in the nominal
subgenus. S. (Pilsbrytyphis) Woodring, 1959,
confined to the middle Miocene of Panama
(Gertman, 1969) also has strange sculpture,
consisting of a wrinkled shell surface. No other
species of Siphonochelus is related, but the
species may be compared with Typhis (Typhina)
bivaricata Verco, 1909. In T. bivaricata the anal
tubes are separated from the varices, which are
foliated and sharp, while rounded and tubercu-
late for the new species ; it has also longer and
more horizontal anal tubes.

Although probably subadult with its 4 teleo-
conch whorls, S. undulatus is definitely distinct
from other Recent and fossil species.

Etymology. Named undulatus for its undu-
lating spiral sculpture.

Subgenus LAEVITYPHIS Cossmann, 1903

Type-species by original désignation : Typhis
muticus J. Sowerby, 1834.

Siphonochelus ( Laevityphis) tillierae
Houart, 1986
Figs 26, 27, 50, 64

Siphonochelus (Laevityphis) tillierae Houart, 1986 :

442, pl. 2, fig. 6.

Material examineo. New Caledonia. “ Vauban "
1978-79 : stn 40. 22"30' S. 166"24' E, 250-350 m 7 June
1979 : 2 shs. (type material).

Biocal : stn CP 75, 22° 19' S, 167“23' E, 825-860 m,
4 September 1985 : 9 shs. Stn DW 79, 20'’40'S,
166"52' E, 1 320-1 380 m. 5 September 1985 : 1 sh -
Stn DW 80. 20°32'S, 166’48‘E. 900-980 m. 5 Sep¬
tember 1985 : 1 sh.

Musorstom 4 : stn DW 161, 18°39'S, 163°U'E,
550 m, 15 September 1985 : 1 sh. Stn DC 168
18"48' S, 163°11' E, 720 m, 16 September 1985 : 1 sh. '

Type locality. New Caledonia, stn 40,
22°30' S, 166"24' E, 250-350 m (“ Vauban ” 1978-
79).

Measurements (holotype). a-b : 5.9 x
2.8 mm. — c : 1.3 x 0.8 mm. - d : 1.3. — e-f :
1.1 x 0.5 mm. G : 70 to 75°.

Description. Shell small and fusiform,
elongate, up to 9.9 mm (Musorstom 4, stn
DW 161). Spire high, consisting of 1.25 to
1.75 protoconch whorls and 4 to 5 rounded
teleoconch whorls. Suture of whorls impressed.
Protoconch rounded and smooth, glossy. Last
whorl bearing 4 strong and somewhat rounded
and shouldered varices. No spiral sculpture.
Axial sculpture consisting of a shallow interva-
rical low ridge, slightly nearer to preceding varix.
A rounded and short anal tube originates near
preceding varix, forming an angle of approxima¬
tely 70 to 75° with the axis of the shell. Other
axial sculpture of line growth striae.

Aperture ovate, columellar lip adhèrent poste-
riorly, olherwise erect. Outer apertural lip erect
and smooth. Siphonal canal short, tubular and
sealed, very slightly bent to the right. Shell
enlirely white.

Discussion. — S. tillierae was originally
compared with Siphonochelus (S.) solus Vella,
1961. 5. (S.) generosus lredale, 1936, S. (Laevi¬
typhis) transcurrens (von M ariens, 1902) and 5.
(Siphonochelus) tubuliger (Thiele, 1925). The
shell of S. tillierae differs from that of S. solus,

Source : MNHN, Paris

MOLLUSCA GASTROPODA : TYPH1NAE

235

S. generosus and S. transcurrens by its smaller
size, form of whorls and form and position of the
anal tubes. From S. tubuliger it difTers by its
more elongate form, its shorter anal tubes and

the angle of these. The aperture of S. tubuliger is
rounded, but ovate in S. tillierae, and the varices
are shorter and much finer.

ACKNOWLEDGEMENTS

I thank Dr P. Bouchet (Mus. natn. Hist.
nat.. Paris) for giving me the opportunity to
study that material and for sem work, and Dr
A. Warén (Nat. Hist. Mus., Stockholm) who
mounted the radulae. Loan of comparative
material was ohtained thanks to most helpful
collaboration of Dr K. Gowlett-Holmes (South

Australian Muséum); Mr B. Marshall (Natl
Mus. New Zealand) and Dr A. Matsukuma
(National Science Mus., Tokyo). I am also most
indebted to Dr P. Bouchet and Prof. E. H.
Vokes (Tulane University) for reading the ma-
nuscript and for their remarks and suggestions.

REFERENCES

Attilio, A. d\ 1975. The typhine shell : theme,
development and élaboration. Feslivus, 6 (9) : 53-60.

Attilio, A. d', 1976. - Recent and fossil Typhinae of
the New World. Feslivus. 7 (5) : 27-30.

Attilio, A. d\ 1979. Interprétation of typhine
morphology with spécial reference to Typhina clarki
(Keen & Campbell, 1964). Feslivus. 11 (11) : 88-93.

Attilio, A. n\ & Hertz, C. M.. 1988. — An
illustrated catalogue of the family Typhidae Cos-
smann, 1903. Feslivus. 20 (supplément) : 1-73, 109
figs.

Fujioka, Y., 1985. Systematic Evaluation of Radu-
lar Characters in Thaidinae (Gastropoda : Muri-
cidae). J. Sci. Hiroshima Univ.. 31 (2) : 235-287,
95 figs.

Gertman, R. L„ 1969. Cenozoic Typhinae (Mol-
lusca : Gastropoda) of the western Atlantic région :
Tulane Stud. Geol. Paleoni.. 7 (4) : 143-191, 8 pis.

Houart. R.. 1983. Three new tropical Muricacean
species (Gastropoda : Muricidae). Venus, Jap. J.
Malac.. 42 (1) : 26-33.

Houart, R.. 1986. Mollusca Gastropoda : Note-
worthy Muricidae from the Pacific Océan with
description of seven new species. ht : Résultats des
Campagnes Musorstom I et 11. Philippines (1976,
1980), volume 2 (13). Mém. Mus. nam. Hist. mil.
Paris , (A). 133 : 427-455.

Houart, R.. 1987a. Description of three new

muricid gastropods from the South-Western Pacific
Océan with comments on new geographical data.
Bull. Mus. nam. Hist. mu. Paris. (4) 8, A, (4) : 757-
767.

Houart, R.. 1987b. Description of four new

species of Muricidae (Mollusca : Gastropoda) from

New Caledonia. Venus. Jap. J. Malac., 46 (4) : 202-
210 .

Houart, R., 1990. — New Taxa and new records of
Indo-Pacific species of Murex and Haustellum (Gas¬
tropoda, Muricidae. Muricinae). Bull. Mus. nam.
Hist. mu. Paris. (4) 12. A, (2) : 329-347.

Iredale, T., 1936. — Australian molluscan notes 2.
Rec. Ausi. Mus.. 19 : 267-340, pis 20-24.

Kaicher, S. D., 1978. — Card catalogue of world¬
wide shells, Muricidae III. Privately publ. St. Peter-
sburg. Fia.

Keen, A. M., 1944. - Catalogue and révision of the
gastropod subfamily Typhinae. J. Paleontology. 18
(1) : 50-72, 20 figs.

Keen, A. M.. & Campbell, G. B.. 1964. Ten new
species of Typhinae (Gastropoda : Muricidae).
Veliger. 7 (1) : 46-57. pis 8-11.

Kool, S. P., 1987. — Significance of radular charac¬
ters in reconstruction of thaidid phylogeny (Neoga-
stropoda : Muricacea). Nautilus. 1Ô1 (3) : 117-132.

Radwin, G., & d'Attilio. A., 1976. Murex shells
of the world. an illustrated guide to the Muricidae :
1-284, 32 pis. Stanford.

Richer de Forges, B., 1990. Les campagnes
d’exploration de la faune bathyalc dans la zone
économique de la Nouvelle-Calédonie. In A.
Crosnier (cd.). Résultats des Campagnes Musors¬
tom. volume 6 (1). Mém. Mus. nam. Hist. nat..
Paris. (A) 145 : 9-54.

Vella, P„ 1961. - Australasian Typhinae (Gastro¬
poda) with notes on the subfamily. J. Palaeomo-
logy, 4 (3) : 362-391. pis 46-47.

Source : MNHN, Paris

Figs 10-18. 10. T y phi s (Typhina) imperialis Keen & Campbell. 1964. New Calcdonia. Musorstom 4 : stn DW 222. mnhn

13.5 mm. II. Typhis /Typliina) carolinae Houart, 1987, New Calcdonia. holotypc mnhn, 20.5 mm. 12. Typhis
( Typhina) virginiae Houart. 1986. New Calcdonia. hololype mnhn. 7.8 mm. 13. Typhis (Talityphis) neocaledonicus
Houarl. 1987. New Calcdonia. holotypc mnhn. 18 mm. 14. Siplionochelus (S.) pavlova (Ircdale, 1936), New
Calcdonia. Musorstom 4 : stn CC 247. mnhn, 10 mm. 15-16. Siplionochelus (S.) angustus sp. nov., holotypc mnhn
6.7 mm. 17-18. Siphonochelus (S.) boucheti sp. nov., holotypc mnhn. 10.1 mm.

Source : MNHN, Paris

Figs 19-27. 19-20. Siphonochelus (S.) saltantis sp. nov., holotype mnhn. 9.2 mm. 21-23. Siphonochelus (S.) unicornis

sp. nov. : 21. New Calcdonia. Biocal : stn DW 46. paratypc mnhn, 9.1 mm. : 22-23. Holotype mnhn. 12 mm. 24-25.
Siphonochelus (? Siphonochelus) undulatus sp. nov.. holotype mnhn. 5.9 mm. 26-27. Siphonochelus (Laevitypliis)
tillierae Houart, 1986. : 26. Holotype mnhn, 5.9 mm.; 27. New Calcdonia. Biocal : stn DW 80. mnhn, 8 mm.

Source : MNHN, Paris

Figs 28-39. 28-29. Siphonochelus (S.) nipponensis Keen & Campbell, 1964. Japan. nsmt 44065, 7.5 mm

30. Siphonochelus (S.) japonicus (A. Adams, 1863), Japan, nsmt 44066. 6.6 mm. 31. Siphonochelus (? Siphonochelus )
syringianus (Hedley. 1903). South Queensland, Auslralia. R. Houart coll.. 8.8 mm. 32-33. Siphonochelus IS ) solus
Vclla. 1961, New Zcaland, nmnz 66725, 9 mm. 34. Siphonochelus (S.) erythrostigma Keen & Campbell, j964 South
Queensland, Australia. R. Houart coll., 10 mm. 35. Siphonochelus (S.) pavlova (Ircdalc. 1936). South Queensland
Australia, R. Houart coll., 15.2 mm. 36-37. Siphonochelus /S.) generosus Ircdalc, 1936, syntype ams C60688
11.8 mm. 38-39. Typhis (Typhina) bivaricata Vcrco, 1909, holotypc sam D1348I, 6 mm.

Source : MNHN, Paris

Figs 40-50. Profiles of aperlurc and last varix of Typhinae (scale lines : 1 mm). 40. Typhis (Typhina/ imperialis,
Musorstom 4 : stn DW 222, mnhn. 41. Typhis (Typhina) virginiae, holotypc mniin 42 Typhis ( Typhinai
carolinae. Lagon : sln 603, mnhn. - 43. Siphonochelus (S.) anguslus , holotypc mnhn. 44. Typhis iTalilyphis)
neoeakdonicus, holotypc mniin. 45. Siphonochelus (S.) pavlova, Musorstom 4 : sln CC 247, mnhn. 46.
Siphonochelus (S.) sahantis, holotype mniin. 47. Siphonochelus 'S.) bouchai, paratype MNHN. 48. Siphonochelus
(S.) unicornis , Biocal : stn DW 46. paratype mnhn. 49. Siphonochelus (? Siphonochelusi undulaïus. holotype mnhn.

50. Siphonochelus (Luevityphis) tillierae. holotype mnhn.

Source : MNHN, Paris

Figs 51-59. — Profiles of apcrlurc and last varix of Typhinae (scalc fines : 1 mm). — 51. Siphonochelus (S.) japonicus, nsmt
44066. — 52. Siplionochelus (S.) nipponensis , nsmt 44067. 53. Siplionochelus (S.) generosus, syntype ams c 60688.

54. Siplionochelus (S.) soins, nmnz m 66725. — 55. Siplionochelus <?Siphonochelus) syringianus. R. HouartcoII. 56.
Siplionochelus (S.) pavlova. R. HouartcoII. —57. Siphonochelus (S.) erythrostigma, R. HouartcoII. 58. Typhis
(Typhina) bivaricata, holotype sam d 13481. 59. Operculum of Typhis (Typliina) carolinae. paralype mnhn.

Source : MNHN, Paris

V

Typhis (Typhina) carolinae . paratype mniin.
(S.) pavlova, " Vauban " : stn 40, mnhn.
Siphonochelus (S.l angusius, hololype mnhn.
phonochelus ( Siphonochelus ) unicorn
mnhn. 69. Siphonochelus (? Sipho

mniin. 71. Siphonochelus (S.) nipponen .

73. Siphonochelus (S.l japonicus. nsmt 44066.
75. Siphonochelus (S.) solus. nmnz m 66725.

60. Typhis (Typhina) imperialis. BlOCAi. : stn DW 64, mniin. 61.
62. Typhis (Typhina) virginiae. holotype mniin. 63. Siphonochelus
64. Siphonochelus (Laevilyphis) tillierae, holotype mnhn. 65.
66. Typhis ( Talilyphis) neocaledonicus, holotype mniin. 67. Si-
Musorstom 4 : sin CC 246, mnhn. 68. Siphonochelus (S.) boucheli, paratype

•helus) undulatus, holotype mniin. 70. Siphonochelus (S. > saltantis, hololype

44065. 72. Siphonochelus ( S.l generosus. syntype a

74. Siphonochelus (? Siphonochelus) syringianus. R. Houart coll.
76. Siphonochelus (S.l erylhrosligma , R. Houart coll.

Source : MNHN, Paris

Source : MNHN, Paris

5ULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSULTATS DES CAMPAGNES MUSORSTOM. VOLUME 7

RÉSUI

10

Mollusca Gastropoda : Columbariform
Gastropods of New Caledonia

M. G. HARASEWYCH

Smithsonian Institution
National Muséum of Natural History
Department of Invertebrate Zoology
Washington, DC 20560 U.S.A.

ABSTRACT

A survey of the deep-water malacofauna of New Caledo¬
nia has brought to light two specics rcferable to the subfamily
Columbariinac (Gastropoda: Turbincllidac). Coluzea faceta
sp. nov. is described from oIT the Isle of Pines at depths of
385-500 m. Addilional specimens of Coluzea pinicola Dar-
ragh. 1987. previously described from off the Isle of Pines.
serve as the basis for the description of the new genus

Fustifusus. Serratifusus virginiae sp. nov. and Serratifusus
lineaius sp. nov., two Recent species of the columbariform
genus Serratifusus Darragh. 1969. previously known only
from deep-water fossil deposits of Mioccne age, are also
described. On the basis of anatomical and radular data,
Serratifusus is transferred from the Columbariinac to the
family Buccinidac.

RÉSUMÉ

Mollusca Gastropoda : Gastéropodes columbariformes de
Nouvelle-Calédonie.

Au cours des campagnes d'exploration de la faune pro¬
fonde de Nouvelle-Calédonie, deux espèces de la sous-famille
Columbariinae (Gastropoda : Turbinellidae) ont été décou¬
vertes. Coluzea faceta sp. nov. est décrite du large de l'île des
Pins entre 385 et 550 m. Coluzea pinicola Darragh, 1987,

également décrite de l'île des Pins, a été récoltée vivante et
devient l'espèce type du nouveau genre Fustifusus. Le genre
Serratifusus Darragh. 1969 n'était jusqu'ici connu que de
dépôts miocènes en faciès profond : deux espèces actuelles, S.
virginiae sp. nov. et S. lineaius sp. nov.. sont maintenant
décrites de Nouvelle-Calédonie. Sur la base des caractères
anatomiques et de la radula. le genre Serratifusus est transféré
des Columbariinae à la famille des Buccinidae.

Harasf.wych, M. G.. 1991. Mollusca Gastropoda : Columbariform Gastropods of New Caledonia. In : A. Crosnier & P. Bouchet
( eds.). Résultats des Campagnes MUSORSTOM. Volume 7. Mêm. Mus. nam. Hist. nul . (A). 150 : 243-259. Paris isbn : 2-85653-180-6.

Publié le 20 mars 1991.

Source : MNHN, Paris

244

M.G. HARASEWYCH

INTRODUCTION

The Columbariinae is the most speciose of the
subfamilies of Turbinellidae, with nearly 50
Recent species inhabiting bathyal depths along
continental margins. predominantly in tropical
and temperate latitudes (Harasewych, 1986).
These animais inhabit soft substrates and feed on
tube-dwelling polychaete worms (Harasewych,
1983). Earliest fossil représentatives occur in the
Late Cretaceous (Maestrichtian) deposits of Eu¬
rope (Darragh, 1969). During the Paleogene,
the Columbariinae were represented in shallow
water (upper continental shelf) faunas ranging
from western North America, throughout Eu¬
rope, to New Zealand (Darragh, 1969). The few
Neogene records are limited to deeper-water
(outer continental shelf — upper continental
slope) fossil deposits of southeastem Australia
and New Zealand (Darragh, 1969; Finlay,
1930), dating the shift in their habitat from
subtidal to bathyal depths. Diverse columbariine
faunas hâve long been known to occur in the
western Atlantic (Clench, 1944, 1959; Bayer,
1971; Harasewych, 1983), western Indian

(Martens, 1901; Tomlin, 1928; Barnard, 1959),
and western Pacific (Habe, 1979; Darragh,
1987; Powell, 1979) Océans. More recently,
columbariines hâve been discovered along conti¬
nental margins of the eastern Pacific (McLean &
Andrade. 1982) and eastern Indian (Harasewy¬
ch, 1986) Océans.

Dr Philippe Bouchet, Curator of Marine
Mollusca, Muséum national d'Histoire naturelle,
Paris, has kindly made available specimens of
columbariform gastropods dredged and trawled
in the upper bathyal zone off New Caledonia as
part of an ongoing faunal sampling program.
The purpose of this paper is to report on this
material, which contained: a new species of
Coluzea; adult specimens and anatomical mate¬
rial of the recently described Coluzea pinicola
Darragh, 1987, that serve as the basis for a
description of a new genus; and two new Recent
species referable to the genus Serratifusus , pre-
viously known only from Early to Middle Mio¬
cène horizons (Darragh, 1969, 1985).

MATERIALS AND METHODS

The présent study is based primarily on speci¬
mens collected by the vessels R.V. " Vauban"
and R.V. “ Jean-Charcot " during Biocal, Mu-
sorstom 4, Lagon, Chalcal 2, and Smib 4
cruises. For a narrative of the deep-sea cruises
and list of stations, see Richer de Forges (1990).

Where sufficient specimens were available,
material for anatomical studies was obtained by
fracturing the shells in a vice, removing larger
fragments with forceps, and dissolving remaining
shell with 10% hydrochloric acid (HCl). Soft
parts were rinsed in distilled water and transfer-
red to 70% éthanol for dissection. Protoconchs
and radulae were cleaned in dilute Sodium
hypochlorite solution (1% NaOCl) prior to exa¬

mination with sem. AU scanning électron micro-
graphs were taken using a Hitachi S-570 sem.

Repositories of examined specimens are indi-
cated by the following abbreviations:
ams Australian Muséum, Sydney
bm(nii)g British Muséum of Natural History,
Geology Department, London
DMNil Delaware Muséum of Natural History,
Wilmington

mnhn Muséum national d'Histoire naturelle,
Paris

nmnz National Muséum of New Zealand,
Wellington

usnm National Muséum of Natural History,
Smithsonian Institution, Washington, DC

Source : MNHN, Paris

MOLLUSCA GASTROPODA : COLUMBARIFORM GASTROPODS

245

SYSTEMATIC ACCOUNT

Family TURBINELLIDAE Swainson, 1840
Subfamily COLUMBAR1INAE Tomlin, 1928

Genus COLLIZEA Allan. 1926

Coluzea Allan, 1926: 304. issued separalely December
7, 1926.

Type species: Fusus deMatus Hutton. 1877, by mono-
typy.

Coluzea Finlay, 1926: 407, issued separalely December
23, 1926.

Type species: Fusus spiralis A. Adams, 1856, by
original désignation.

The morphological similarities belween lhe
généra Coluzea, and Fulgurofusus Grabau, 1904.
hâve been discussed previously (Finlay, 1930:
267-268; Harasewych, 1983: 5. 1986: 158). Full
generie slatus is provisionally retained for Colu¬
zea until a révision of the subfamily can be

completed. Coluzea first appears in the Eocene
deposits of the Paris and London Basins. and is
known from the Late Oligocène of New Zealand
(Darragh. 1969). As presenlly understood. the
genus appears to be restricted in the Recent
fauna to continental margins in the temperate
and tropical régions of the Indian Océan, to the
eastern margin of the Australian tectonic plate,
and to those adjacent régions of the Pacific
tectonic plate west of the Andésite Line (see
Springer, 1982: fig. 2). A request for a ruling by
the International Commission on Zoological
Nomenclature on the authorship and type spe¬
cies of Coluzea (Beu et ai, 1969) is still pending.

Coluzea faceta sp. nov.
Figs 1-4, 22; Table 1

Material examined. New C'aledonia. “ Vau-
ban", 1978: stn 14. 22°16'S, 167°17'E. 465-495 m,
23-28.05.1978: paratype 10, 49.5 mm, mnhn.

Musorstom 4: stn 239. 22"14.80'S, 167°15.70‘E,
470-475 m, 2.10.85: paratype 9, 58.5 mm. mnhn.

Stn 242, 22°05.80'S, I67°I0.30'E, 500-550 m, 3.10.
1985: holotype 82.6 mm, mnhn. Stn 247,
22°09.00'S, 167°13.30’E. 435-460 m, 4.10.1985: paraty-
pes 1-2, 44.7 mm, 43.7 mm, usnm 860477: paratypes
3-5, 50.0 mm, 39.5 mm, 23.6 mm, mnhn; paratype 6,
38.1 mm. ams. Stn 248, 22°09.50'S, 167‘10.00’E.
380-385 m, 4.10.1985: paratype 7, 57.4 mm, mnhn;
paratype 8, 43.4 mm, nmnz.

Type Material. Holotype, mnhn; paraty¬
pes 1-2, usnm 860477; paratypes 3-5,7, 9, 10,
mnhn; paratype 6, ams; paratype 8, nmnz.

Type Locality. Off southeastern New
Caledonia, 22°05.80’S. 167°10.30'E, in 500-550
m. [Musorstom 4, stn 242],

Description. Shell (Figs 1-2) large (to 83
mm), heavy. fusiform: spire angle 28°-37°: pro-
toconch (Fig. 4) of 1 1/4 whorls, with first whorl
deflected from coiling axis by about 80°; transi¬
tion to teleoconch marked by appearance of
keeled axial ribs. followed within 1 whorl by
spiral cords above and below shell periphery;
teleoconch with up to 10 convex. weakly shoul-
dered whorls; suture adpressed; axial sculpture of

9- 10 hollow ribs on early postnuclear whorls.

10- 11 thick, solid varices with short, open spines
along periphery on sixth and subséquent whorls;
spiral sculpture of 4-5 strong cords between
suture and shoulder. 4-5 between shoulder and
anterior carina. 4- 5 between anterior carina and
base of siphonal canal, 32-38 cords on proximal
2/3 of siphonal canal; 1-4 fine threads between
adjacent spiral cords; second, and occasionally
first spiral cord anterior to periphery particularly

Source : MNHN, Paris

Figs 1-4. Coluzea facela sp. nov. : 1, Hololypc, mnhn, Musorstom 4, stn 242. Apcrlural. latéral and dorsal views. (0.775

x). 2. Paratype 7. mnhn. Musorstom 4. stn 248. Apcrlural view. (0.775 x). 3. Opcrculum of paratype 7. (3.1 x).

4. Protoconch and first teleoconch whorl. paratype 5. mnhn, Musorstom 4. stn 247. (24.8 x).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : COLUMBARIFORM GASTROPODS

Figs 5-9. Fustifusus pinicola (Darragh. 1987) : 5. Juvénile spccimen of lhe samc size as the holotype. mniin. Smib 4. stn
8. Apcrtural and dorsal views. (1.86x). 6, Adull specimen. mnhn. Biocal, sln 42. Apcrtural. latéral and dorsal views.

( 1.24 x). - 7, Sub-adult specimen, mnhn, "Vauban". 1978, stn 4. Apertural view. (1.24 x). 8, Opcrculum of spccimen

in figure 2. (4.34 x). 9, Protoconch and first telcoconch whorl, "Vauban". 1978 stn 4. (24.8 x).

Source : MNHN, Paris

248

M. G. HARASEWYCH

pronounced, giving posterior portions of axial
ribs angular, banded appearance; aperture ovate;
outer lip thick, furrowed beneath periphery,
inner lip smooth; siphonal canal 1/2 shell length,
axial, stout, slightly sinuate distally; shell color
white; periostracum thin, yellowish, consisting of
sériés of fine, axial, blade-like lamellae; oper-
culum (Fig. 3) thin, amber-colored, strongly
ovate, with sharply tapered terminal nucléus.

Table I. Coluzea face la sp. nov.

characlers. Linear measuremenls in ram. N

Charactcr
Shell length
Aperture length
Aperture length

Shell length
Siphonal canal length
Siphonal canal length

Shell length
No. whorls teleoconch
No. varices, body whorl
Spire angle

Measurements of shell
10 .

X Range SD
47.9 23.6-82.6 14.7
7.4 3.6-12.6 2.2

0.16 0.14-0.17 0.01
23.7 15.6-40.4 7.3

0.49 0.41-0.55 0.05
7.8 7.0-10.0 1.2
11.2 10-13 0.9
32.8° 28°-37° 2.5°

Animai. One male (paratype 9) and two female
(holotype and paratype 7) specimens were dis-
sected. Mantle cavity spanned about 3/4 whorl,
kidney, 1 /3 whorl, digestive gland not preserved;
animal uniform tan in color; foot rectangular
(L/W = 2.0); tentacles short; eyes absent or
unpigmented; mantle edge thickened; osphra¬
dium short, broad (L/W = 3.0). with 65 leafiets
above and 50 below axis; ctenidium 1.2 limes as
wide, 2.5 times as long as osphradium, with
about 450 leafiets; hypobranchial gland transver-
sely pleated; rectum long, narrow; rectal gland
along anterior 1/3 of rectum; anus attached to
roof of mantle cavity, with short ventral papilla;
extended proboscis 1.1 times shell length; radula
(Fig. 22) short (2 mm), narrow (110 pm), with
105-133 rows of teeth (n = 3); latéral teeth
monocuspid, with broad basal plate; rachidian
teeth tricuspid, with laterally expanded basal
plates; valve of Leiblein large, completely contai-
ning région of oesophageal torsion; salivary
glands asymmetrical, adhering to each other and
to oesophagus; salivary ducts enter oesophagus
anterior to valve of Leiblein, becoming embed-
ded beneath dorsal folds; gland of Leiblein large,
occupying posterior 1/2 of cephalic sinus, situa-
ted to left and below proboscis sheath; posterior

oesophagus, broad, sacculate along gland of
Leiblein, constricting at rear of cephalic sinus;
stomach simple, U-shaped; rectum broad poste-
riorly, tapering anteriorly; female palliai gono-
duct with long, tall capsule gland; bursa copula-
trix ovate, anterior to capsule gland; male palliai
gonoduct of simple narrow duct descending to
fioor of mantle cavity at mid-length to form
deep, open, muscular groove lined with glandu-
lar tissue, running along inner edge of short,
dorsoventrally-flattened pénis; pénis with glan-
dular pad at distal end, without papilla; nervous
System highly concentrated, with circumoesopha-
geal ganglia fused.

Etymology. — L. facetus, well made, élégant.

Distribution. This species is known from 5
stations, ail off the Southern tip of New Caledo-
nia. The confirmed bathymétrie range is 385-500
m, with a mean station depth of 461.5 m. Living
specimens were taken at ail stations except
" Vauban" 1978, station 14.

Ecology. — Little is known of the habitat of
this species, other than the bathymétrie range.
Nearly ail specimens hâve more than one repai-
red shell break, indicating frequent, unsuccessful
prédation by crabs and/or fish. Ail cases of shell
breakage were limited to two varices or less (<
1/5 whorl), testifying to the effectiveness of the
thickened costae as antipredatory adaptations.
None of the specimens had been drilled.

Comparative Remarks. - Coluzea faceta is
conchologically most similar to Coluzea distepha-
notis (Melvill. 1891), but may be distinguished
from this and the other Recent Western Austra-
lian and New Zealand species of Coluzea by its
heavier shell, coarse spiral sculpture, and its
pronounced and thickened axial costae. The lack
of a canaliculate suture differentiates it from the
Indonesian species C. liriope Harasewych, 1986
as well as from the geographically most proximal
C. bimurata Darragh, 1987, which inhabits shal-
lower depths off Queensland. Coluzea faceta also
resembles C. angularis (Barnard, 1959) from off
South Africa, but differs in having short, open
spines along the peripheral keel, and by the
rounder shape of its aperture. When compared
to the New Zealand species of Coluzea , C. faceta
more closely resembles C. paucispinosa Finlay,
1930 and C. dentala (Hutton, 1877), both from
the Miocene, than any younger species.

Source : MNHN, Paris

MOL1.USCA GASTROPODA : COLUMBARIFORM GASTROPODS

249

Genus FUSTIFUSUS gen. nov.

TypeSpecies. Coluzea pinicola Darragh, 1987:

133-134, figs 4 E,I.

Description. Columbariinae with heavy,
fusiform, high-spired shell of moderate size (to
46 mm); protoconch, glassy, inflated, deflected
from coiling axis by about 60°, first whorl as
large or larger than subséquent whorl; spiral
sculpture predominating in early whorls, forming
suturai canal by fourth or fifth post-nuclear
whorl; axial sculpture increasing in prominence
with increasing shell size, forming pronounced
ribs or varices with open peripheral spines by
seventh post-nuclear whorl: color light brown
with bands of créant or white along periphery, at
juncture of body whorl with siphonal canal, and
along siphonal canal (Fig. 6); basal plate of
rachidian tooth U-shaped (Fig. 24), but not
laterally expanded.

Etymology. Latin fustis, a knobbed stick,
fusus , a spindle.

Comparative Remarks. The globose.
strongly deflected protoconch, pigmented shell.
and weak but distinct raised columellar plate
indicate that this genus is more closely related lo
Columbarium that lo any of the généra or
subgenera in the Coluzea Fulgurofusus lineage.
It differs from most Recent species of Columba¬
rium in retaining the plesiomorphic high spire, in
having broad, thickened varices, and a channeled
suture, as well as in lacking numerous serrated
spiral cords along the siphonal canal. The radula
of the type species is unlike that of any known
columbariine in having two usually well-defined
but smaller cusps on either side of the three large
cusps of the rachidian tooth. The recurved basal
plate lacking laterally-expanded butresses is a
charaeter shared with species of Columbarium
[e.g. Columbarium spinicinclum (von Martens,
1881) see Harasewych, 1983: fig. 11].

Fustifusus pinicola (Darragh, 1987)
Figs 5-9, 23-24: Table 2

Synonymy: Coluzea pinicola Darragh, 1987: 133-134,
figs 4 E.l.

Material examined. New Caledonia: “ Vau-
han 1978: stn 4. 22°17’S. 167°13'E, 400 m. 23-
28.05.1978: 3 specimens 37.0 mm, 26.6 mm, 14.5 mm.
MNIIN. Stn 16, 22°46’S, 167°I2 , E, 390-400 m,
10-15.04.1978: 1 specimen 17.9 mm, mnhn.

Lagon: stn 395, 22°48.2’S, 167°07.6'E. 313 m,
23.01.1985: 1 specimen 31.9 mm. mnhn. — Stn 419,
22°42.3’S, I67°10.5'E, 330 m, 24.01.1985: 1 specimen

23.5 mm, mnhn.

Biocal: stn 38, 22°59.74'S. 167*15.3l'E. 360 m,
30.08.1985: 1 specimen 35.44 mm, mnhn. Stn 42.
22°45.14'S, 167*12.12‘E, 380 m, 30.08.1985: 1 speci¬
men 45.2 mm. mnhn. Stn 44, 22°47.30’S,
167° 14.30'E, 440-450 m. 30.08.1985: 7 specimens 20.8
mm, 19.4 mm, 19.3 mm. 19.2 mm. 19.0 mm. 17.8 mm.

16.6 mm. mnhn.

Musorstom 4: stn 212, 22°47.40’S, I67°10.50'E,
375-380 m, 28.09.1985: 1 specimen 23.1 mm, mnhn.
Stn 226. 22°47.20'S, 167*21.60'E. 395 m, 30.09.1985: 3
specimens 41.2 mm. usnm 860478. 33.5 mm, 8.8 mm.
mnhn. Stn 227, 22°46.00 S. 167°20.00'E. 320 m.

30.09.1985: 1 specimen 20.3 mm, mnhn. — Stn 230,
22°52.50’S, 167*11.80’E. 390-420 m. 30.09.1985: 3
specimens 25.4 mm, 21.1 mm, 15.4 mm, mnhn. Stn
234. 22°15.50’S. 167°08.30’E. 350-365 m. 2.10.1985: I
specimen 36.6 mm, usnm 860479.

Smib 4: stn 5. 22°56.3'S, 167°14.4'E, 398-410 m,
17.09.1986: 1 specimen 20.4 mm. mnhn. Stn 8.
22*53.6'S, 167° 12.5'E. 435-447 m, 18.09.1986: 1 speci¬
men 21.1 mm, mnhn.

ca. 22°50'S, 167°I5'E. 200-400 m: 3 specimens 40.4
mm, 35.2 mm, 24.13 mm. mnhn.

Type Material. Holotype. ams. C82163:
paratype 1. ams. Cl52010.

Type Locality. South of Isle of Pines,
New Caledonia. 22°50’S. 167°35.5'E, in 370 m.
[“ Kimbla " stn K4-71-4],

Description. Shell (Figs 5-7) of medium
size (to 46 mm), moderately heavy, fusiform:
spire angle 28°-33.5°; protoconch (Fig. 9) of 1

Source : MNHN, Paris

250

M. G. HARASEWYCH

whorl, glassy, rotund; first 1/2 whorl deflected
from shell axis by about 60°, larger than subsé¬
quent whorl; transition to teleoconch graduai,
marked by formation of peripheral keel, rounded
at first, becoming progressively more acute;
nodules develop along keel by second postnu-
clear whorl, hecoming short, open spines by
fourlh post-nuclear whorl; teleoconch with up to
7 convex whorls; suture adpressed; spiral sculp¬
ture of 3-4 cords between suture and periphery,
5-6 between periphery and siphonal canal, 20-28
on proximal 2/3 of siphonal canal; adsutural
spiral cord increases in prominence, forming
suturai canal by fourth post-nuclear whorl; axial
sculpture of 8-16 low ribs per whorl, forming
spines and nodes at intersections with peripheral
keel and adjacent spiral cords, respectively;
growth striae fine, sinuate: aperture strongly
ovate to triangular; outer lip thick, glazed,
furrowed beneath peripheral keel; inner lip
smooth, with raised inductural edge extending
length of siphonal canal in adult specimens;
siphonal canal long, straight. stout; base color
ginger to brown, with lighter bands along peri¬
phery and siphonal canal; aperture white, occa-
sionally with brown band along outer lip; perios-
tracum thin, ginger brown, with fine axial
lamellae forming tufts along spiral cords and rim
of suturai canal; operculum (Fig. 8) thin, amber-
colored, rounded posteriorly, sharply tapered
anteriorly, with terminal nucléus.

Table 2. Fustifusus pinicola (Darragh, 1987). Measure-
ments of shell characters. Linear measurements in mm.
n = 10.

X Range SD

35.3 23.1-45.5 6.4

5.6 4.5-6.7 0.8

0.16 0.14-0.19 0.02

16.9 9.4-23.4 4.5

0.50 0.45-0.53 0.02

6.9 6.0-7.3 0.4

10.4 8-13 1.3

31.3° 28°-33.5° 1.5”

Animal : Two dried male specimens were rehy-
drated and dissected. Foot long, narrow, rectan-
gular (l/w = 2.2); tentacles short, blunt, with
large black eyes; mantle cavity narrow, of 1 1/4

whorl; osphradium large, broad, ctenidium equal
in width to osphradium; hypobranchial gland
whitish, globular, 1 1/2 times as wide as cteni¬
dium; pénis short, narrow, blunt, with open
sperm groove; proboscis long, folded within
proboscis sheath; buccal mass minute; radula
(Figs 23, 24) short (980 pm ), narrow ( 60 pm),
consisting of 102-118 rows (n = 2); basal plate of
rachidian tooth U-shaped, with 1-2 short cusps
on either side of 3 long central cusps; latéral
teeth with single, long, scythe- shaped cusps.

Etymology. — Named after the type localily,
the Isle of Pi nés.

Distribution. This species has been collec-
ted at 15 stations, ail off the Isle of Pines. The
confirmed bathymétrie range for live-collected
specimens is 330-440 m. Specimens inhabited by
hermit crabs hâve been taken as shallow as 300
m. The mean station depth for ail records is
373.2 m.

Ecology. — This species inhabits soft subs¬
trates at depths of 330 to 440 meters, and
co-occurs with Serratifusus virginiae sp. nov.,
described helow, at 6 of the 15 stations. Poly-
chaete setae were found in the rectum of one
specimen of Fustifusus pinicola.

Comparative Remarks. — This species was
originally described in the genus Coluzea , based
on the morphology of two immature specimens
(Darragh, 1987). Allhough juvénile specimens
of Fustifusus pinicola resemble New Zealand
Miocene species of Coluzea in having high-spired
shells with strong spiral sculpture and weak
tubercles along the keel, these characters are
probably symplesiomorphic within Columbarii-
nae and therefore not indicative of close phylo-
genetic relationships. The globose protoconch,
weakly raised columellar plate, and diffuse
brown coloration présent in this species are
characters that occur in most western Pacific
Columbarium species, but not in species Coluzea
or Fulgurofusus. The thickened varices that ge-
nerally do not appear until the seventh post-
nuclear whorl also occur in Columbarium nata-
lense Tomlin, 1928, a species that inhabits
shallower (90-200 m) depths off southeastern
Africa.

Character
Shell lenglh
Aperture length

Shell length
Siphonal canal length
Siphonal canal length

Shell length
No. whorls teleoconch
No. varices, body whorl
Spire angle

Source : MNHN, Paris

MOLLUSCA GASTROPODA : COLUMBAR1FORM GASTROPODS

251

Family BUCCINIDAE Rafinesque, 1815
Subfamily BUCC1N1NAE Rafinesque, 1815

Genus SERRATIFUSUS Darragh, 1969

Serratifusus Darragh. 1969: 89

Type Species. Fusus craspedotus Tate, 1888.
by original désignation.

In his révision of the family Columbariidae,
Darragh (1969: 90) proposed the genus Serra¬
tifusus for a group of closely related fossil species
that: hâve prominent axial sculpture on the final
whorl of the protoconch, hâve axial ribs on early
teleoconch whorls, hâve a pronounced peripheral
keel with open spines, and that lack a curved
lamella on the columellar lip. He recognized that
this genus differed from Columbarium, but retai-
ned it in the then family Columbariidae on the
basis of the conchological resemblance to Colu-
zea. Serratifusus was believed to be reslricted to
Early to Middle Miocene deposits of southeas-
tern Australia and Tasmania (Darragh. 1969.
1985).

A newly discovered Recent species, described
below, closely resembles Serratifusus craspedotus
(Tate, 1888), the type species of Serratifusus, in
features of shell and protoconch. and exhibits ail
the diagnostic characters of this genus. Thus, the
stratigraphie range of Serratifusus is extended to
the Recent. An examination of the anatomical
organization of the Recent species reveals that
Serratifusus is more properly assigned to the
family Buccinidae. Tricuspid rachidian leeth with
a rectangular basal plates, tricuspid latéral teeth,
a narrow, sacculate gland of Leiblein, a closed

vas deferens, a pénis with a closed sperm duct,
and the lack of a rectal gland are among the
characters that support the transfer of the genus
to the Buccinidae, and suggest that Serratifusus
is closely related to Penion Fischer. 1884 (see
Ponder. 1973).

In discussing the type species of Serratifusus.
(as Fusus craspedotus) Harris (1897: 54. 135)
commented on its resemblance to “ certain forms
of Columbarium ", but assigned il to the family
Fasciolariidae on the basis of protoconch mor-
phology. Serratifusus youngi (Chapman, 1922)
was provisionally included in Serratifusus. but
may prove to be referable to Fusinus (Darragh.
1969: 92). The eight remaining species originally
included in Serratifusus (Darragh. 1969) most
likely represent a monophyletic assemblage wi-
thin the Buccinidae.

Serratifusus appears most closely related to
Penion based on general conchological similarity
as well as nearly identical anatomical organiza¬
tion and radular morphology (Ponder. 1973).
Serratifusus may be distinguished from Penion,
however. by its smaller. thinner shell. longer
siphonal canal, and by its sharply angled peri-
phery with prominent radial spines. Serratifusus
and Penion are found togelher in Eariy to
Middle Miocene beds (Darragh. 1989. personal
communication). In the Recent fauna. Serratifu¬
sus is restricted to bathyal depths, while Penion
occurs in shallower waters along the continental
slope.

Serratifusus virginiae sp. nov.
Figs 10-12, 17. 19. 25; Table 3.

Material examined. New Caledonia. " Vau-
ban" , 1978: stn 16. 22“46’S. 167°12'E, 390-400 m.
10-15.04.1978: paratype 2, 40.5 mm. mnhn.

Musorstom 4: stn 212, 22°47.40'S,167°10.50’E, 375-
380 m, 28.09.1985: holotype. 40.7 mm, mnhn; pa¬
ratype 1. 45.2 mm, USNM 860480. Stn 226,

22°47.20’S. 167*21.60’E. 395 m. 30.09.1985: paratypes
3-4. 34.9 mm. 28.9 mm. mnhn. Stn 227. 22°46.00’S,
167*20.00’E. 320 m. 30.09.1985: paratype 5. 35.6 mm.
ams; paratype 6. 32.8 mm. mnhn. — Stn 234.
22*15.50’S. 167*08.30’E, 350-365 m. 2.10.1985: paraty¬
pes 7-8, 39.0 mm, 30.7 mm, mnhn.

Source : MNHN, Paris

252

M.G. HARASEWYCH

Fig. 13. Serrati/usus craspcdolus (Taie. 1888). BM(nh) G.38761-5. Muddy Creek near Hamilton. Victoria. Australia.
Balcombian (Middlc Miocène). Apertural and dorsal vicws (1.24 x).

Figs 10-12. Serratifusus virginiae sp. nov. : 10. Holotype, mnhn. Musorstom 4. sln 212. Apertural. latéral and dorsal views.
(1.24 x). Il, Paratype I. usnm 860480 . Musorstom 4. stn 212. Apertural and dorsal views. (1.24 x). 12. Paratype

7. mnhn, Musorstom 4. stn 234. Apertural view. (1.24 x).

Source : MNHN, Paris

MOLLUSCA GASTROPODA : COLUMBAR1FORM GASTROPODS

253

Figs. 14-16, Serratifusus lineatus sp. nov. : 14. Holotypc, mnhn. Musorstom 4, stn 181. Apertural, latéral and dorsal views.
(1.24 x). 15. Paratype I. usnm 860843. Musorstom 4. stn 181. apertural and dorsal views. (1.24 x). 16. Operculum

of holotype of Serratifusus lineatus sp. nov. (3.1 x).

Fig. 17 — Serratifusus virginiae sp. nov. : Holotypc. mnhn. Musorstom 4. stn 212. Operculum.

Source : MNHN, Paris

254

M.G. HARASEWYCH

Chalcal 2: stn 81, 23°19.60'S, 168°03.40'E. 311 m,
31.10.1986: paratype 10. 38.2 mm, nmnz; paratypes
11-12, 19.4 mm, 18.5 mm, usnm 860481. Stn 82,
23°13.68'S, 168°04.27'E, 304 m. 31.10.1986: paratypes
13-14, 31.0 mm, 30.7 mm. mnhn.

From fishing boat, ca. 22°50 , S, 167“15'E, 200-400
m: paratype 9, 31.7 mm, mnhn.

Type Material. — Holotype, mnhn; paratype
1. usnm 860480; paratypes 2-4, mnhn; paratype
5, ams; paratypes 7-9. mnhn; paratype 10. nmnz;
paratypes 11-12, usnm 860481; paratype 13-14,
mnhn.

Type i.ocality. West of the Isle of Pines,
New Caledonia, 22°47.40'S, 167°10.50'E, in 375-
380 m. [Musorstom 4, stn 212].

Description. Shell (Figs 10-12) large (to 46
mm), thin, fusiform; spire angle 45-55°; proto-
conch (Fig. 19) of 1 1/2 inflated whorls, first
whorl deviated from coiling axis by 45°; transi¬
tion to teleoconch marked by smooth axial ribs
with knobs along periphery becoming gradually
more pronounced, forming short, broad, open
spines by third postnuclear whorl; teleoconch
with up to 6 inflated, sharply shouldered whorls;
suture adpressed; spiral sculpture weak, area
between suture and peripheral keel smooth, with
weak threads along spines; body whorl below
periphery with 19-21 cords; cords equally broad
posteriorly, gradually differentiating into alter-
nating broad and narrow cords anteriorly; 36-38
alternating broad and narrow spiral cords on
proximal 2/3 of siphonal canal; axial sculpture of
6-8 ribs on early whorls, forming broad spines on
later whorls; growth striae very fine, strongly
sinuate; aperture subtriangular; outer lip furro-
wed beneath peripheral keel; inner lip smooth;
siphonal canal long, slightly sinuate; shell color
light tan to ginger, with white blotches on spines
and reddish brown blotches mosl strongly pro¬
nounced between suture and periphery and along
siphonal canal; aperture white to light tan;
periostracum thin, amber-colored, axially lamel-
late; operculum (Fig. 17), broadly oval, with
terminal nucléus and médian ridge.

Anima !: One male (paratype 5) and two female
(holotype and paratype 1) specimens were dis-
sected. Soft parts span 2 1/2 whorls, mantle
cavity 1/2 whorl, kidney 1/4 whorl, digestive
gland 1 1/2 whorls; animal yellowish tan, with

Table 3. Serratifusus virginiae sp. nov. Measuremcnls of
shell characiers. Linear measuremcnls in mm. N = 9.

Characler
Shell lenglh
Aperlure lenglh
Aperture lcnelh
Shell lenglh
Siphonal canal lenglh
Siphonal canal lcnelh

Shell lenglh
No. whorls teleoconch
No. varices, body whorl
Spire angle

X Range SD

34.8 17.3-45.2 7.8

8.8 6.1-11.8 2.7

0.25 0.19-0.32 0.04

15.4 8.6-21.3 4.2

0.44 0.36-0.53 0.07

5.0 3.5-5.75 0.6

10.2 8-12 l.l

49.6° 45.5°-55.0“ 3.2°

fine black spots on head, foot; foot broad (l/w
= 1.2); tentacles short, broad; eyes large, black;
mantle edge thin, smooth; osphradium long,
broad (L/W = 3.2), with 97 leaflets above and
78 leaflets below axis; ctenidium equally long, 1
3/4 times as wide as osphradium, with about 280
leaflets; hypobranchial gland narrower than cte¬
nidium; rectum broad posteriorly, tapering ante¬
riorly; rectal gland absent; anus pendant; exten-
ded proboscis about 0.4 times shell length; radula
(Fig. 25) short (2 mm), narrow (80 pm), of
90-112 rows of teeth (n = 3); latéral teeth
tricuspid, outermost cusp longest, emanating
from near center of basal plate, central cusp
shortest, thinnest; rachidian teeth tricuspid, cen¬
tral cusp longest, basal plate deep, rectangular;
valve of Leiblein large, anterior to région of
esophageal torsion; salivary ducts paired, sur-
rounding valve of Leiblein and nerve ring; gland
of Leiblein, thin, brown, silualed along left side
of retracted proboscis; stomach simple, tubular;
female palliai gonoduct comprised of albumen
gland, situated along right side of kidney, large
ingesting gland, long, narrow capsule gland, and
large, muscular, anteriorly silualed bursa copu-
latrix; male palliai gonoduct consists of closed
muscular duct of uniform diameter, glandular
along posterior 3/4 of length, leading from rear
of mantle cavity to base of pénis; pénis short ( 1 /2
mantle cavity length). distally broad, dorsoven-
trally compressed, with short papilla along trun-
cated distal outer edge; nervous System highly
concentrated.

Etymolouy. This species honors Ms Virgi¬
nie Héros for her work in processing much of

Source : MNHN, Paris

MOLLUSCA GASTROPODA : COLUMBARIFORM GASTROPODS

255

20

Figs. 18-20. Protoconch and carly tclcoconch whorls of spccics of Serralifusus. Ail vicws 15.5 x : 18, Serratifusus craspedotus
( laie, 1888), bm(nh) G.38761-5. muddy Creek near Hamilton, Victoria. Australia. Balcombian (Middle Mioccnc). -
19, Serralifusus virginiae sp. nov., Musorstom 4, stn 227. 20, Serralifusus lineatus sp. nov„ Musorstom 4, stn 181.

Source : MNHN, Paris

256

M.G. HARASEWYCH

Radulae. Orienlcd with anterior end toward lop of the page. Ail scalc bars = 25 gm.

Fig. 21. Coluzea spiralis (A. Adams. 1856) dmnh 48393. Off Kawau ld.. North Island. New Zealand.

Fig. 22. Coluzea faceta sp. nov. Holotypc, mnhn, Musorstom 4, sln 242.

Fig. 23-24. Fustifusus pinicola (Darragh. 1987). usnm 860479. Musorstom 4. stn 234;

23. Radula in rctractcd orientation, with latéral teclh folded over rachidian teeth.

24. Radula in extended, rasping orientation, with latéral teeth unfolded.

Fig. 25. Serratifusus virginiae sp. nov. Holotype, mniin, Musorstom 4. stn 212.

Fig. 26. Serratifusus lineatus sp. nov. Hololype. mnhn, Musorstom 4. stn 181.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : COLUMBARIFORM GASTROPODS

257

the deep-water expédition material collected off
New Caledonia.

Distribution. — This species has been collec¬
ted at 8 stations, ail off the Isle of Pines. The
confirmed bathymétrie range is 311-395 m for
live-collected specimens. The mean station depth
for ail specimens is 343.4 m.

Ecology. This species inhabits soft subs¬
trates at depths of 300 to 390 meters, and
co-occurs with Fustifusus pinicola (Darragh,
1987) at 6 of 8 stations. No identifiable gut
contents were found in any of the three speci¬
mens examined.

Comparative Remarks. - The inclusion of
this new species in the genus Serratifusus is based

on its distinctive protoconch as well as features
of the sculpture, aperture and peripheral keel
(Darragh, 1969; 69). Adult specimens of Serra-
tifusus virginiae closely resemble Serratifusus
craspedotus (Tate, 1888) (Fig. 13), but differ in
being higher-spired and narrower, in having
fewer, heavier and longer spines, and in lacking
pronounced spiral cords between the suture and
shell periphery. Sub-adult specimens (Fig. 12) of
5. virginiae hâve proportionally longer siphonal
canals, and thus may bear a doser resemblance
to S. stellatus Darragh, 1969. The latter species
differs from S. virginiae in having a higher spire,
a thicker, stouter siphonal canal, and two or
three scaly lirae along the siphonal canal.

Serratifusus lineatus sp. nov.
Figs. 14-16, 20, 26; Table 4.

Material examined. New Caledonia. Musors-
tom 4: stn 156. 18°54.00'S. 163°18.80'E, 525 m.
15.09.1985: 1 specimen (fragment) 37.7 mm, mnhn. —
Stn 164, 18°33.20'S, 163°13.00’E, 255 m, 16.09.1985: 1
specimen 24.5 mm. mnhn. Stn 181, 18°57.20'S,
163°22.40'E, 355 m. 18.09.1985: holotype, 35.6 mm,
mnhn; paratypes 1-3, 31.0 mm, 30.4 mm. 27.7 mm,
usnm 860843; paratypes 4-18, 40.9 14.3 mm. mnhn;

paratype 19, 35.3 mm. ams; paratype 20, 25.4 mm,
nmnz. — Stn 184. 19°04.00'S. I63°27.50’E. 260 m,
18.09.1985: paratypes 21-31, 43.0 — 10.4 mm, mnhn.

Stn 195, 18°54.80'S, 163°22.20'E, 470 m,

19.09.1985: paratypes 32-40. 32.2 - 19.1 mm. mnhn.

Stn 196, 18°55.00’S, 163°23.70’E, 460 m,

20.09.1985: paratypes 41-46.30.7 17.9 mm. mnhn.

Type Material. Holotype, mnhn; paraty¬

pes 1-3, usnm 860843; paratypes 4-18, mnhn;
paratype 19, ams; paratype 20, nmnz; paratypes
21-46, mnhn.

Type Locality. Western end of Grand

Passage, off northwestern New Caledonia,
18°57.20'S. 163°22.40'E, in 350 m. [Musorstom
4, stn 181],

Description. Shell (Figs 14-15) large (to 42
mm), thick, fusiform; spire angle 48°-55°; proto¬
conch (Fig. 20) conical, of 1 1/2 whorls, first
whorl deviated from coiling axis by 50°; transi¬
tion to teleoconch marked by axial ribs with

pronounced knobs along periphery that form
short, broad, solid spines by third postnuclear
whorl; teleoconch with up to 7 inflated, strongly
shouldered whorls; suture adpressed; spiral
sculpture almost enlirely lacking, limited to faint
threads that correspond to spiral Unes of dark
brown pigment; axial sculpture of 6-8 broad fiat
solid spines along the shoulder; growth striae
fine, sinuate; aperture ovate; outer lip smooth, or
with 8-12 weak denticles in large specimens;
inner lip smooth; siphonal canal slightly longer
than aperture, sinuate; shell color white to light
tan, with parallel, dark brown bands that may be
solid or interrupted, 3 between suture and peri¬
phery, 1 along periphery. 8-9 on body whorl, and
7-8 on proximal 1/2 of siphonal canal; periostra-
cum very thin, yellowish; operculum (Fig. 16),
narrow, roughly semicircular, with terminal nu¬
cléus and curved médian ridge on outer surface.

Animal : One male (holotype) and one female
(paratype 1) specimen were dissected. Tissues
span 2 3/4 whorls, mantle cavity 1/2 whorl,
kidney 1/4 whorl, digestive gland 1 3/4 whorls;
animal cream colored, with fine black spots on
head, foot; foot broad (l/w = 1.4); tentacles
short, broad; eyes large, black; mantle edge thin.
smooth; osphradium (l/w = 2.2) as wide and
2/3 as long as ctenidium. with 110 leafiets above

Source : MNHN, Paris

258

M. G. HARASEWYCH

Table 4. Serralifusus lineatus sp. nov. Measurements of
shell characters. Linear measurements in mm. N = 10.

Character
Shell length
Aperture length
Aperiure length

Shell length
Siphonal canal length
Siphonal canal length

Shell length
No. whorls teleoconch
No. varices, body whorl
Spire angle

X Range SD
34.6 27.7-41.1 3.6
10.9 9.3-12.2 0.8

0.32 0.30-0.34 0.02
11.1 8.7-13.5 1.3

0.32 0.30-0.33 0.01
6.0 5.5-6.5 0.2
7.4 7-8 0.5
51.8° 48.0°-55.0° 2.2°

and 92 leaflets below axis; ctenidium with about
350 leaflets; hypobranchial gland narrow, pen¬
dant, glandular; rectum broad posteriorly, tape-
ring anteriorly, detaching from roof of mantle
cavity just posterior to anus; rectal gland absent;
extended proboscis about 0.5 shell length; radula
(Fig. 26) short (3.5 mm), narrow (180 pm), of
98-124 rows of teeth (n = 3); latéral teeth tricus-
pid, outermost cusps longer than basal plates,
emanating from near center of basal plates,
central cusps narrowest; rachidian teeth tricus-
pid, cusps of equal length, basal plates deep,
rectangular; valve of Leiblein large, anterior to
région of esophageal torsion; salivary glands
paired, surrounding valve of Leiblein and nerve
ring; gland of Leiblein, thin, brown, running
along left side of proboscis sheath; stomach
simple, tubular, with two widely separated ducts
to digestive gland; female palliai oviduct consis-
ting of albumen gland along right wall of kidney.

ingesting gland, elongate capsule gland, and
large, pyriform bursa copulatrix situated anterior
to capsule gland; male palliai gonoduct consists
of closed muscular duct of constant diameter,
leading from rear of mantle cavity to base of
pénis; pénis long (2/3 mantle cavity length),
dorsoventrally compressed, tapering distally,
with short papilla along truncated outer edge.

Etymology. — Latin lineatus, marked with
Unes.

Distribution. — This species is known only
from six stations, ail off the western end of the
Grand Passage. The confirmed bathymétrie
range for live-collected specimens is 260-470 m,
although fragments and hermit-crab occupied
specimens were taken from 255 m to 525 m. The
mean station depth for ail specimens is 387.5 m,
for live-collected specimens 386.3 m.

Ecology. — Little is known of the ecology of
this species other than it inhabits rubble bottoms
at depths of 260-470 m. No identifiable gut
contents were found in any of the three speci¬
mens examined.

Comparative Remarks. — The shorter sipho¬
nal canal, smaller but thicker spines along the
shoulder, lack of spiral sculpture, and the cha-
racteristic coloration consisting of parallel brown
spiral Unes serve to distinguish this species from
Serralifusus virginiae, its only presently known
Recent congener.

ACKNOWLEDGEMENTS

1 am grateful to Thomas A. Darragh,
Muséum of Victoria, and Bruce A. Marshall,
National Muséum of New Zealand, for their

reviews of an earlier draft of this manuscript.
Their comments and suggestions hâve substan-
tially improved this paper.

REFERENCES

Allen, R. S., 1926. — The geology and paleontology
of the Lower Waihao Basin, South Canterbury,
New Zealand. Trans. N. Z. Inst.. 57: 265-309.

Barnard, K. H.. 1959. - Contributions to the

knowledge of South African marine mollusca. Part
II. Gastropoda: Prosobranchia: Rachiglossa. Ann
S. Afr. Mus.. 45(1): 1-237.

Source : MNHN, Paris

MOLLUSCA GASTROPODA : COLUMBAR1FORM GASTROPODS

259

Bayer, F. M., 1971. New and unusual mollusks
eollected by R/V John Elliot Pillsbury and R/V
Gerda in the tropical western Atlantic. Bull. mur.
Sci., 21(1): 111-236.

Beu, A. G., Dell, R. K., Fleming, C. A.,
Marwick, J.. Maxwell. P. A., Ponder, W. F.,
& Powell, A. VV. B., 1969. — Requests for rulings
on Works on New Zealand Mollusca by R. S. Allan
and H. J. Finlay. Z.N. (S.) 1868. Bull. zool. Nom., 26
(1): 42-50.

Chapmann, F., 1922. — New or little—known fossils
in the National Muséum. Part XXVI: SomeTertiary
mollusca. Proc. Roy. Soc. Viet.. 35(1): 1-18, pl. 1-3.

Clench. W. J., 1944. — The genus Columbarium in
the western Atlantic. Johnsonia, 1(15): 1-4.

Clench. W. J.. 1959. The genus Columbarium in
the western Atlantic. Johnsonia, 3(39): 330-331.

Darragh, T. A.. 1969. — A révision of the family
Columbariidae (Mollusca: Gastropoda). Proc. R.
Soc. Vici.. 83 (1): 63-119, pis 2-6.

Darragh. T. A.. 1985. — Molluscan biogeography
and biostratigraphy of the Tertiary of southeastern
Australia. Alcheringa. 9: 83-116.

Darragh, T. A., 1987. Columbariinae (Gastro¬
poda; Turbinellidae) from Queensland, Australia.
Proc. R. Soc. Vict., 99 (4): 127-134.

Finlay, H. J., 1926. — Further commentary on New
Zealand Molluscan systematics. Trans. N. Z. Inst..
57: 320-485, pis 18-23.

Finlay. H. J., 1930. — Révision of the New Zealand
Shells referred to Fusinus. Trans. N. Z. Inst., 61:
259-270, pis 47-49.

Grahau, A. W., 1904. Phylogeny of Fusus and its
allies. Smithson. mise. Colins.. 44(1417): 1-192, pis
1-18.

Habe, T.. 1979. The western Pacific columbariid
species. Venus. 38(1): 1-4.

Harasewych, M.G., 1983. A review of the
Columbariinae (Gastropoda: Turbinellidae) of the
western Atlantic with notes on the anatomy and

systematic relationship of the subfamily. Nemouria.
27: 1-42.

Harasewych, M.G., 1986. — The Columbariinae
(Gastropoda: Turbinellidae) of the eastern Indian
Océan. J. malac. Soc. Au si.. 1 (3-4): 155-170.

Harris, G. F., 1897. Catalogue of the Tertiary
Mollusca. Part 1. Australasia. British Muséum (Nat.
Hist.) London: xxvt + 407 p„ 8 pis.

Martens, E. von, 1901. Einige ncue Meer-
Conchylien von der deutschen Tiefsee-Expedition.
Sber. Ges. nalurf. Freunde. Berlin: 14-26.

McLean. J. H. & Andrade, H., 1982. Large
archibenthal gastropods of central Chile: Collec¬
tions from an expédition of the R/V Anton Bruun
and the Chilean Shrimp Fishery. Comr. Sci. nat.
Hist. Mus. L.A. County. 342: 1-20.

Ponder, W.F.. 1973. A review of the Australian
species of Penion Fischer (Neogastropoda: Buccini-
dae). J. malac. Soc. Ausl., 2 (4): 401-428.

Powell, A. W. B., 1979. - New Zealand Mollusca.
Marine, Land and Freshwater Shells. Collins, Auck¬
land: xiv + 500 p., 82 pis, 119 figs.

Richer de Forges, B., 1990. — Les campagnes
d'exploration de la faune bathyale dans la zone
économique de la Nouvelle-Calédonie. Explorations
for bathyal fauna in the New Caledonian économie
zone. In : A. Crosnier (ed.). Résultats des Campa¬
gnes Musorstom, Volume 6. Mcm. Mus. nain. Hist.
nat., (A) 145 : 9-54.

Springer, V. G., 1982. — Pacific plate biogeography.
with spécial reference to shorefishes. Smithson.
Conirib. Zool.. 367: 1- 182.

Swainson, W., 1840. — A treatise on malacology; or
the natural classification of shells and shellfish. A.
Spottiswoode, London: viii + 419 p.

Taxe, R.. 1888. The gastropods of the older
Tertiary of Australia. Part I. Trans. R. Soc. S. Ausl.
10: 91-176. pl. 1-13.

Tomlin, J. R.. 1928. Reports on the marine
mollusca in the collections of the South African
Muséum. Ann. S. Afr. Mus.. 25(2): 313-335.

Source : MNHN, Paris

Source : MNHN, Paris

ACHEVÉ D’iMPRIMER
EN MARS 1991
SUR LES PRESSES

L’IMPRIMERIE F. PAILLART
À ABBEVILLE

DÉPÔT LÊCAL : 1" TRIMESTRE 1991
IMP. 7685. DISTRIBUÉ LE 20 MARS 1991

Source : MNHN, Paris

Source : MNHN, Paris

Source : MNHN, Paris

22 MARS 1991

Source : MNHN, Paris

DERNIERS TITRES PARUS
RECENTLY PUBLISHED MEMOIRS

Série A (Zoologie) :

Tome 149 : Jean Chazeau & Simon Tillier (eds), 1991 — Zoologia Neocaledonica. Volume 2. 358 pp. (ISBN
2-85653-179-2) 400 FF.

Tome 148 : Loïc Matile, 1990 — Recherches sur la systématique et l'évolution des Keroplatidae (Diptera,
Mycetophiloidea). 682 pp. (ISBN 2-85653-173-3) 650 FF.

Tome 147 : Nicole Berti (ed.), 1990 - Miscellanées sur les Staphylins. 222 pp. (ISBN 2-85653-158-X) 300 FF.

Tome 146 : Christian Erard, 1990 — Écologie et comportement des gobe-mouches (Aves : Muscicapinae,
Platysteirinae. Monarchinae ) du Nord-Est du Gabon. Volume 2 : Organisation sociale et écologie de la
reproduction des Muscicapinae. 234 pp. (ISBN 2-85653-178-4) 320 FF.

Tome 145 : Alain Crosnier (ed.). 1990 Résultats des Campagnes Musorstom. Volume 6. 388 pp. (ISBN 2-
85653-171-7) 440 FF.

Série B (Botanique) :

Tome 32 : Claudine Friedberg, 1990 Le savoir botanique des Bunaq. Percevoir et classer dans le Haut
Lamaknen (Timor. Indonésie). 304 pp. (ISBN 2-85653-177-6) 350 FF.

Tome 31 : Odile Poncy, 1985 — Le genre Inga (Légumineuses. Mimosoideae) en Guyane française.

Systématique, Morphologie des formes juvéniles. Ecologie. 124 pp. (ISBN 2-85653-135-0) 210 FF.
Tome 30 : Lucile Allorge, 1985 Monographie des Apocynacées — Tabernaemontanoïdées américaines.
216 pp. (ISBN 2-85653-132-6) 280 FF.

Tome 29 : Monique Keddam-Mai pi.anche, 1985 — Le Pollen et les stomates des Gardéniées (Rubiacées) du
Gabon. Morphologie et tendances évolutives. 109 pp. (ISBN 2-85653-131-C-8) 220 FF.

Tome 28 : Marie-France Roquebert. 1981 Analyse des phénomènes pariétaux au cours de la conidiogenèse
chez quelques Champignons microscopiques. 79 pp. (ISBN 2-85653-116-4) 130 FF.

Série C (Sciences de la Terre) :

Tome 56 : Jean-Paul Saint Martin, 1990 Les formations récifales coralliennes du Miocène supérieur
d'Algérie et du Maroc. 373 pp. (ISBN 2-85653-170-9) 392 FF.

Tome 55 : Georges Busson (ed.), 1988 Évaporites et hydrocarbures. 144 pp. (ISBN 2-85653-155-5)
180 FF.

Tome 54 : Monette Véran, 1988 — Les éléments accessoires de l'arc hyoïdien des poissons téléostomes (Acan-
thodiens et Osteichthyens) fossiles et actuels. 114 pp. (ISBN 2-85653-154-7) 150 FF.

Tome 53 : Donald E. Russell, Jean-Pierre Santoro and Denise Sigogneau-Russell, 1988 Teeth
Revisited : Proceedings of the Vllth International Symposium on Dental Morphology. 462 pp. (ISBN
2-85653-148-2) 625 FF.

Tome 10 : Jacques Roger, 1962 (Réimpression/Reprint 1988) - Buffon. Les Époques de la nature. Édi¬
tion critique. 344 pp. (ISBN 2-85653-160-1) 100 FF.

Prix hors taxe, valides jusqu'à décembre 1991. Frais de port en sus. Vente en France : TVA 2,10%.
Prices in French Francs are valid until December 1991. Postage not included.

Source : MNHN, Paris

La zone économique maritime de la Nouvelle-Calédonie occupe 1,5 millions de kilomètres carrés
dans le sud-ouest de l’Océan Pacifique. Depuis 1984, les navires de recherche français ont effectué une série
de campagnes océanographiques pour l’exploration de la faune profonde de ce secteur. Plusieurs centaines
de dragages et chalutages, jusqu’à 3700 m de profondeur, ont permis la découverte d’une faune d’une
grande nouveauté, dont l’étude a été entreprise par les chercheurs de nombreux pays.

Le présent volume, le septième de la série des « Résultats des Campagnes Musorstom », est
consacré aux Mollusques bathyaux et abyssaux de Nouvelle-Calédonie. L’extrême originalité et la
remarquable diversité de cette faune apparaissent nettement à travers les dix articles : ce ne sont en effet
pas moins d’une famille et une sous-famille, 5 genres et 90 espèces nouvelles qui sont décrits dans ce
volume. Trois genres de Gastéropodes, jusqu’ici connus uniquement à l’état fossile d’Australie et d’Asie du
Sud-Est, sont signalés pour la première fois dans la faune actuelle.

Alain Crosnier, qui est Directeur de Recherche à I’orstom, coordonne l’étude de la faune profonde
de Nouvelle-Calédonie et la publication des résultats scientifiques. Il a été secondé pour ce volume par
Philippe Bouchet, Sous-Directeur au Muséum national d’Histoire naturelle.

ÉDITIONS
DU MUSÉUM
38,

RUE GEOFFROY
SAINT-HILAIRE
75005 PARIS
ISBN 2-85653-180-6
ISSN 0078-9747

PRIX : 357 FF TTC (France)
350 FF (Étranger)

Source : MNHM, Paris